THE GENUS OPISTHORCHIS

BY

ARSHAG KILLIJIAN SEUERIAN

A. B. Ripon College, 1918

THESIS

Submitted in Partial Fulfillment of the Requirements for the

Degree of

MASTER OF ARTS

IN ZOOLOGY

IN

THE GRADUATE SCHOOL

OF THE

UNIVERSITY OF ILLINOIS

1919

UNIVERSITY OF ILLINOIS

THE GRADUATE SCHOOL

I HEREBY RECOMMEND THAT THE THESIS PREPARED UNDER MY

SUPERVISION RY Arahag Killijian Seuerian

ENTITLED The Genua jflpiathorch ia

BE ACCEPTED AS FULFILLING THIS PART OF THE REQUIREMENTS FOR

THE DEGREE OF Maater of Arta _

Recommendation concurred in*

Committee

on

Final Examination*

Required for doctor's degree but not for master's

UIUC

TABLE OF CONTENTS

I. Introduction

II. Hiatory

III. Synopsis of the species once named in the genus

Opda*horchia> and there division into three Genera:

Opisthorchis , Amphimerus and the genus Parapisthorchis

IV. Discussion

V. Description of Opisthorchia viverrini Var .Chiengmai

1. Material and Technique

3. History

3. General description of the form

4. The excretory system

5. The reproductive system

6. The alimentayr system.

7. The body layer a, glands and parenchymas

8. The nervous system

9. Discussion.

VI. Bibliography

VII. Explanation of Plates

INTRODUCTION

This investigation haa been carried on under the direc-

tion of Dr. H. B. Ward with material furnished by him. The bottle

containing the material was labeled Bpisthorchis viverrini (?$.

The specimens were collected in 1915 by Dr. C. W. Maaon from the

liver of a prisoner who died in the Chiengmai jail, Siam. Hundreds

of thewe forms were sent to Dr. Ward.

Leiper 1915 made a provisional diagnosis of this form and

called it Opisthorohis viverrini.

The object of the present paper is two-fold: first to give

a detailed description of this form; seoond, to bring all data

available on the genus Opistorchis in a single work for a ready

reference in the study of the problem of species and varieties

among the closely related species now existing.

Teohnique. - The specimens were somewhat poorly preserved.

The material was stained in to to with Erlich's acid hematoxylin,

borax carmine, and fuchsin. Cross, frontal and sagittal sections

were made and stained with dilute Delafield's hematoxylin and with

Erlich's aoid hematoxylin countered stained with eosln. A great

number of specimens were examined for marked variations. Twenty

specimens were however selected at random for general measurements.

Digitized by the Internet Archive

in 2013

http://archive.org/details/genusopisthorchiOOseue

- 2 -

II. HISTORY

Loose (189^) made a few genera out of various species

classified under the genus Opisthorchie . He placed these genera

( Holometra, Metorchis, Clonorchis and Opisthorchis) under the

subfamily Opisthorchi inae . The specific diagnosis of this no

subfamily Opisthorchiinae Looss (1899:563) is as follows:

Fasciolidae of medium size with a slender elongated body,

anterior end noticeably tapering. Suckers near one another and

usually not well developed. Pharynx present. Intestinal crura

long and simple, short and slender oesophagus. Exoretory kladder

Y-shaped with a long S-shaped branch winding between the testes

and with short side branches. Genital pore commonly in front of

the acetabulum. Copulation organs absent. Testes near the hinder

end more or less obliquely placed. Ovary is in front of the tes-

tes. Laurer's canal present. Seminal receptacle very strongly

developed. Uterus loops in front of the testes. Vitellaria

moderately developed, lateral to the intestinal crura.

Found in the liver of mammals, birds and reptiles.

The above diagnosis and the three new genara belonging to

the new subfamily, in its modified form has been accepted by all

parasitologists.

Campula was originally proposed for distomes with a wavy

intestine by Cobbold (1885). Stiles and Haesall did not find

sufficient ground for the separation. "Opisthorchis was based

upon the topography of the genital organs, Campula upon the crook-

ed intestine. The two genera are, however, hardly to be separat-

ed at present, since Campula agrees in topography with Opisthorchis'

»

- 3 -

(Paras. 1:85). Later Cobbold himself admitted the identity of

the two species ( jfc885:39). Looss expresses the same opinion as

Stiles and Hassall (1889)

Key to the genera of the subfamily Opisthorchiinae Looss (1899)

1. (2) Vitellaria lateral to intestinal crura 3

2. (5) Vitellaria anterior to intestinal crura, lateral

to oesophagus; uterine coils fill entire body post-

erior to acetabulum - - - - - - HOLOMETRA

3. (2) Vitellaria not extending anteriorly beyond the

acetabulum ----------------- 4

4. Vitellaria extending beyond the acetabulum

METORCHIS

5. (6) Testes dendritic ------ - - - - -7

6. Testes not dendritic OPISTHOPCHIS

7. Testes dendritic CLONORCHIS

The diagnosis of the genus Opisthorchis Blanchard and the

modifications made by Looss are placed below for comparison.

Opisthorchis R. Blanchard (1895:217). - Medium and large

forms. Oesophagus long or short, occasionally wanting. Intestin-

al branches not ramified. Genital orifice in front of acetabulum.

No cirrus pouch. Vitelline glands generally do not extend behind

the testes. Ovary situated posterior to uterine coils. Testes

simple, globular and ramified.

Genus Opisthorchis Blanchard in partim.- Body transparent,

often very strongly attenuated, posterior end bread; tapering

toward the anterior end. Skin smooth with or without spines.

- 4 -

Testes more or less strongly lobed or ramified, lie obliquely one

in front of the other. Ovary lobed or simple. Uterus loops lying

in front of the ovary and not usually reaching out over the in-

testinal crura. Vitellaria does not extend forward beyond the

ventral acetabulum •

The type of the genus is Opisthorchis felineus.

From the above diagnosis of the genus Opisthorchis I leave

out species once reported as Opisthorchis which do not agree to

the modified genus Opisthorchis. For example D. albidue Braun

(1893) , according to the later modification of 0pi8thorchie

Blanchard, falls into the genus Metorchis Looss (1899). 0. exigua

Mflhl. (1898) comes under Holometra. 0. sinensis Cobbold (1815)

under Clonorchis. In the gemiB Metorchis are 0. crassiuscula

Rud:.1809 var. (?~sp . nov. ? ) Janus. 0. crassiuscula janus

Kowal. (1898h), 0. crassiuscula Looss (1899b:678), 0. xanthosoma

Crepl? Wag. , var

.

(?) campascua.

The species belonging to the genus Opisthorchis at present

are as follows:

0. felinius Riv. 1884

0. tenuicollis Rud. 1819

0. viverrini Poirier 1886

0. longiseAmus Linet. 1883

0. simulans Looss 1896

0. geminus Looss 1896

0. piscicola Odhner 1902

0. obsequens Nicoll 1915

o. entzi w. Ratz 1900

- 5 -

0. interruptue Eraun 1901

0. lancea Diesing 1850

0. noverca Braun 1902

0. pseudofelinaua Ward 1895

0. epiolosus Stiles and Hassan 1896

0. ovalis Earker 1911

0. oaninus Barker 1911

- 6 -

III. SYNOPSIS OF THE SPECIES ONCE NAMED IN THE GENUS

OPISTORCHIS: AND THERE DIVISION INTO THREE GENERA: CPISTH*

ORCHIS, AMPHIMERUSi AND PARAPISTHORCHIS'.

(Ward 1903 and Stephens 1916)

The type species designated for the genus Opisthorchis is

0. felineus Rivolta (1884)* Gurlt named the fluke Distoma oonus

in 1831, but Creplln had already used the name for another species

in 1825. Von Siebold's Distoma lanceolatuw 1836, was preoooupied

by the gemie known as Dicrocoelium dindriticum. Distoma sibirioum

proposed by Winogradoff 1892 cannot be accepted by law of prior-

ity and in fact it is a variety of 0. felineus Riv. Rivolta gave

D. felineus in 1884 which holds* D. tenuieolle, Muhl . 1896.

Diagnosis - Body much flattened, yellowish in color and al-

most transparent in living specimens. The sides of the body are

parallel save in the region anterior to the acetabulum which is

conical. There is a shallow constriction at the level of the

ventral sucker noticeable in fresh and contracted specimens. The

posterior end is either pointed or rounded off. The length and

breadth vary according to the contraction, being usually 8 to

11mm by 1.5 to 2 mm. The suckers are about one-fifth to one-sixth

of the body length distant from each other, and about equal in

size. 0.23 to 0.25 mm. The oesophagus is very short. The intest-

inal crura reach almost to the posterior border and are often fil-

led with blood. The excretory pore is at the posterior extremity,

and the excretory bladder forks in front of the anterior testis.

Anterior testis four lobed, posterior five lobed, and occupy the

posterior fourth of the body one placed obliquely in front of the

other. Ovary median, transverse, simple or slightly lobed.

- 7 -

Smaller than the pear or retort-shaped receptaculum seminalis.

Laurer's canal present. Uterus in the median field and entirely

within the intestinal crura. The vitellaria occupy the fairly

broad lateral areas in about the central third of the body, be-

ginning behind the ventral sucker and terminating at about the

level of the ovary. The acinic are small and are arranged in groups

of seven to eight, separated by interstices. The genital pore

is immediately preace tabular . Egg* oval with distinct operculum

at the pointed pole, 0.030 by 0.011 mm.

Habitat - Gall duct and gall bladder of cats, dogs, glutton,

fox and man.

Distribution - Europe , (Germany , France, Holland, Scandinavia,

Hungary and Russia) and Siberia.

Opistorchis viverrini Poirier 1886

Body transparent, lanceolate, 6mm long and 2 mm broad . The

dorsal surface a little concave the ventral surface flat. Suckers

small and equal in size, 0.23 mm. Ventral sucker at the beginning

of the anterior third. The distance between the two suckers 1.8 mm.

Genital orifice immediately above the ventral sucker, "phe

length of the pharynx 0.1mm immediately poster i»e to the oral

sucker, which has a wide opening. Oesophagus is relatively short

0.3 mm. The intestinal forks are simple and with a nearly con-

stant deameter C.lmm and terminate blindly at the extreme post-

erior end.

The ovary is situated in front of the testes and la multi-

lobed. It extends transversely, having two well developed lobes

on the legt and two smaller ones on the right. The shell glands

in this case present a peculiar aspect. They are not compact

- 8 -

but small unicellular isolated glande scattered in the parenchyma

around the beginning of the uterus.

The racemose vitelline glands are situated laterally in the

median region of the body and in front they dissappear before

reaching the level of the ventral sucker. The single transverse

vitelline duct has the vitelline reservoir on the left of the

shell gland. Laurer's oanal very distinct at the base of which

the pear shaped receptaculum seminal is is situated just outside

of the shell gland. The uterus is very strongly coiled and sit-

uated in the region of the body between the ventral sucker and

the ovary.

The eggs are small, ovoid, 0.026 mm long and 0.013 mm wide.

The male organs are composed of two small four lobed testes;

which are situated at the posterior of the body. The vas defeaens

is a very narrow and long duct. It forms a coiled pouch at the

right of the middle line of the body and extends to the genital

cloaca.

The excretory apparatus is not clearly distinguished. The

excretory bladder is quite voluminous and extends from the ex-

ternal pore at the posterior end to the ovary.

These were collected from the bile duct of Felix viverrinius

which died in the cattle shed of a Museum in France. ( Poirier

1886:6)

Opisthorchis geminus Loose (1896:50)

The body flattened, elongated, tapering anteriorly, increas-

ing in size posteriorly. Total length 7 to 8 mm, in Circus

aeruginosis 10 to 12.5 by 1 to 1.7 mm with a cephalad region

0.2 mm. The greatest width 1.3 mm. Oral sucker quite mall 0.17

- 9 -

mm in diameter. The oral opening subterminal. Acetabulum smaller

than the anterior sucker, situated at the second fourth of the

body

.

The integument very delicate, smooth and without any spicules

or spinelwts.

Digestive system - The mouth is followed by the small phar-

ynx measuring not more than 0.1 mm in diameter followed by the

slender owsophague 0.25mm long. The intestinal crura are not

thicker than the oesophagus and extend almost to the posterior

end bending slightly toward the median pland lying at the sides

of the excretory bladder at the extreme posterior end.

The Y-shaped excretory system curved slightly forming an

"S" between the testes. The short branches at the anterior end

do not extend over the intestine.

Genital system - The genital orifice is marked by a small

opening immediately anterior to the acetabulum, gopulatdry

organs not noteced. The two testes are not situated exactly

at the middle line of the body. The anterior one is placed some-

what to the left, the posterior to the right. The anterior testis

four, the posterior usually five lbbed; their greatest diameter

not exceeding 0.6 mm. Each gives a small vas efferens uniting at

the middle of the body forcing the vas deferens which is very

large in diameterO.l mm, and very long. After many transverse

loops which do not reach the intestinal branches laterally it

opens into the genital sinus.

The ovary usually trilobed, situated in front of the testes,

is generally smaller than the testes and nearly in the middle

line of the bftdy. Laurer's canal is prominently curved and filled

with spermatozoa. In the interior of the germ duct cilia sse

- 10 -

present. The vitellaria occupy the lateral portion of the body

outside of the intestinal branches. The vitelline glands begin

about the anterior third or middle of the length of the body and

extend not further than the posterior end of the ovary. The traas«

verse ducts consequently are posterior to the vitellaria. The

course in fact is not perpendicular to the longitudinal axis of

the body but extends somewhat posteriorly. The much coiled uterus

does not extend over the intestinal branches. It opens into the

genital sinus.

The egga are oval in shape with a lid, 0.02 mm long and

0.01 mm wide. Perfectly developed embryos are seen in the metra-

term region.

Habitat - In the biliary duct of Milvus parasiticus, Anas

boschas fera, Oircua aeruginosus.

Distribution - Egyp$.

Opisthorohis simulans Looss (1896:52)

Three adult specimens and one young form found by Loess in

the biliary duct of Pernis apivorue captured near Alexandria.

The worms were located near the liver in the biliary passage.

The specific diagnosis - Body flat, elongated, anterior end

less tapering, posterior end slightly tapering. The length is

7 to 23 mm, the greatest width across the middle of the body 1.5mm.

The narrow anterior end 0.6 mm. The anterior sucker surrounding

the oral opening is terminal, it is relatively large, strong and

measures 0.5 mm in diameter, the acetabulum on the other hand is

quite small and dees not measure more than 0.2 mm in diameter,

it is situated nearly at the lefel of the anterior third of the

body

.

- 11 -

The integument is smooth without spines or spinelete.

Digestive system - The pharynx well developed 0.3 mm in dia-

meter nearly in contact with the befuroation of the crura so that

no oesophagus is present. The onura run to the posterior end of

the body widening a little at the extremity.

The nervous system - The nervous system is similat to khat

of the other members of this genus.

The excretory system - The excretory sfttem is Y-shaped. The

unpaired bladder winds between the testes and has two short

branches, into which opens a system of vessels at the sides of

the ovary

.

The genital system. - The genital orifice is at the anterior

border of the acetabulum and opens into the genital sinus. The

testes have a median position are spherical or elongated, 0.6 mm

in diameter. The aasa efferentia start from the lateral anterior

borders of the testes and join 0.8 mm posterior to the middle of

the acetabulum. By this union they form in this species the

single vas deferens 0.1 mm in maximum diameter which opens into

the male orifice after having a winding course.

The ovary is quite small, aobed (follicular) and is situated

in front of the testes at the left and adjoining the middle plane

of the body. The receptaoulum seminis is small and olub shaped.

Laurer's canal is quite short and a little swollen at the region

which follows the receptaculum seminis. At the anterior portion

of the oviduot as in other parts cilia «re present. The vitel-

laria occupy the space at the sides of the body outside of the

intestinal branches. They are very much ramified and extend to

the anterior border of the anterior testis. The uterus occupies

- 12 -

the anterior portion of the female ganital organs, -phe loope

wind transversely and do not extend over the intestinal crura,

it opens into the common genital sinus.

The eggs are larger tfcan in 0. geminus and measure 0.038 mm

in length by 0.018 mm in width. The operculum is a little thinner

The shell in thin but less transparent on account of the brown

color. Eggs in the region of the metraterm have not developed

embryos

.

Habitat - Liver of Anas penelope, Fulix cristata, Cireus

aeruginosus and Pernis apivorus.

Distribution - Egypt.

Opisthorchis obsequens Hicoll (1914:106)

The body is very much flattened with crenate edges. The

anterior end is considerably attenuated but the posterior end is

rounded. A small knob-like tip projects however from the anterior

end. The adult specimens measure 2.6 to 5.0 mm. The maximum

breadth which occurs in the post- acetabular region, varies

from 0.8 to 1.2 mm. The average specimen measures 3.7 by 1. mm.

There are no spines. The oral sucker is almost terminal and

measures 0.16 by 0.20 mm on the average. Its transverse diameter

varies from 0.15 to 0.24 mm. The ventral sucker measures 0.26mm

and is situated 1.24 mm from the anterior end, which is exactly

one- third of the body length.

There is no prepharynx. The pharynx has a diameter of 0.12 mm

and is slightly longer than broad. The oesophagus is about

0.19 mm in length with a range of 0.13 to 0.29 mm. The intestin-

al diverticula are sinuous and narrow with crenate walls, espec-

ially on their inner side. They extend to within 0,2 mm of the

- 13 -

posterior end of the body. Their ends are turned in towards the

middle line.

The excretory vesicle has the usual sinuous configuration

observed in the other species of the genus. The posterior curve

may be either towards the right or to the left. The bifurcation

occurs just behind the ovary and the limbs are short.

The genital aperture is median and immediately in frcmt of

the ventral sucker. The vesicula seminalis is short and highly

convoluted. It extends beyond the ventral sucker for a distance

about 0.3 mm. The testes are very much lobed, almost dendritic.

There are usually five lobes on the anterior testis and four on

the posterior; but this is not invariable. They are more or less

tandem, while there is always a strong tendency to obliquity,

the anterior testis being usually displaced towards the left side •

Amphitypy occurs frequently. The posterior testicular space

measures 0.34 mm which is about one-eleventh of the body length.

The ovary lies in front of the testes and is almost median.

It is excessively fcobed, there being usually three lobes directed

backwards. It is much smaller than either of the testes. The

receptaculum seminis is of median size, pear-shaped and lies as

a rule on the right. The yolk glands extend from the level of

the middle of the ventral sucker to the level of the anterior

testis and usually not beyond. They lie entirely to the outer

side of the intestinal diverticula and consist of eight groups

of follicles on either side. The transverse yolk ducts meet

at a very obtuse angle in froat of the ovary. The uterus entire-

fills the space "between the ovary and the ventral suoker. It

forms from the fourteen to sixteen transverse convolutions which

occasionally touch, but do not overlap, the intestinal diverticula

- 14 -

The egg measures 0.025 to 0.030 by 0.014 to 0.017 mm the average

being 0.028 by 0.0155 mm.

Habitat - Liver of Hieracidea berigora (brown Hawk) and

Hieracidea orientalis.

Distribution - North Queensland.

Opiethorohis entace Von Ratz, (1900: 532-534)

"Body flattened dorso-ventrally , lanceolate in shape and

transparent; anterior portion whitish and posterior brownish yel-

low; anterior end tapering and constricted at the lefel of the

acetabulum forming a neck region anterior to the acetabulum; post-

terior portion of the body broader and the end rounded; length 12

to 13.2 mm; breadth at the widest part 1 to 1.2 mm. Oral sucker

0.25 mm in diameter; acetabulum 2.38 mm posterior to the oral

sucker and a little larger, measuring 0.31 ram in diameter.

Pharynx oval or heart shaped and followed by a short o»sophagus

and forked intestinal tract, the crura ending in blind enlargments

in the posterior end of the body. Cirrus and cirrus sac absent.

Testes lie in the posterior half of the body, the one behind the

other, 4 to 5 lobed. Vas efferens from each testes unite to

form the vas deferns which passes forward to the genital pore

situated just anterior to th* acetabulum. Ovary in the middle

of the body and slightly lobed, the retort shaped receptaculum

seminis and tubular Laurer*s canal just posterior to the ovary.

Vitellaria lateral to the intestinal caeca and extend from near

the acetabulum to the ovary. Acini of the vitelline glands some-

what indefinite being crowded together in spaces. The vitelline

glands extend farther oephalad on the right side than ofl the left.

The two vitelline ducts, one from each side, lead posteriorly to

- 15 -

the ovary where they unite. Uterine coils well developed and fill

the body between the intestinal crura and the ovary and the

acetabulum, but the coils do not extend over the intestinal crura.

The shell gland is distinct and just anterior to the ovary. Sggs

oval, more pointed at the anterior end where the operculum with

an opercular rim is found. They measure 0.0242 mm by 0.0132 to

0.0154 mm".

Habitat - Blue Heron ( Ardea purpurea L)

Distribution - Hungary.

Opisthorchie tenuicollis Rud. (1819:4261

Specific diagnosis - Body lanceolate, flat anterior end

markedly tapering and thin, posterior end bluntly pointed; body

6.5 Co 10.12 mm by 1.13 mm. Oral sucker 0.134 by 0.152 mm. Acet-

abulum 0.219 mm in diameter, situated between the first and second

fourth of the body. Cuticula without spines. Ovary oval, smooth •

Receptaculum seminis well developed. Laurer's canal and shell

gland not mentioned. Testes in the posterior fourth of the

body. Four to five lobed. Vitellaria in one region lateral to

the intestinal caeca. Vitelline glands begin just posterior to

the acetabulum and extend to the ovary. Uterus protfuBely devel-

oped. Eggs elliptical. 0.0228 by 0.014 mm.

Habitat - Reported from the domestic cat? dcg? and man?

Gulo boreal is, Pooa barbata, Halichorus grypus,

Erignathus barbatus, Dephinus phocaena, Phocaena phoc-

aena.

Distribution. - Europe.

Opisthorchis piscicola Odhner (1902:152)

Body about 8 to 4 mm long. Greatest breadth about 0,5 mm

Ilea in the region of the middle of the body. From that the worm

tapers strongly toward the anterior, much less strongly toward ttee

posterior. The forward end is rounded. The first part of the

posterior end behind the testes is abruptly pointed. Mouth open-

ing circular 0.19 to 0.23 mm in diameter, ventral acetabulum

transveraly oval. 0.25 by 0.3 mm. Situated in the end of the

first body fifth. The body is armed with small broad spines. The

pharynx is 0.08 mm in diameter. Testes strongly lobed lying in

the median line. Ovary is more or less crenate. Genital glands

immediately posterior to one another. The vitellaria extends

from the posterior end of the ovary anteriad to the middle third

of the body. Ripe eggs 0.021 mm long and 0.011 mm broad.

Habitat - Gall bladder of Nile fish (Gymnarohus niloticus)

Distribution - White Nile. Omdurman.

Opisthorchis pseudofelineus Ward (1895:152-158)

Body elongated, transparent, flat; length 5 to 22 mm; breadth

1 to 2.5 mm. Anterior end conical; posterior end rounded. Oral

sucker 0.24 to 0.541 mm. Ventral sucker 0.24 to 0.28 mm in dia-

meter, situated about one-fourth of the total length of the body

from the oral sucker. Pharynx 0.12 to 0.25 mm. long by 0.14 to

0128 mm wideand neat to the oral suoker, owsophagus short 0.04

to 0.38 mm in length; branches of the intestine extend nearly to

the posterior end of the body . Two lobed testes in the poeterior

third or fourth of the body, the poeterior lying obliquely caudad

to the anterior. Anterior orbicular, oval or elliptical in shape;

margin smooth, wayy or 2 to 5 lobed. Ovary elliptical, orbicular

or oval in shape with margin smooth, wavy or lobed, situated a

little in front of the testes. Receptaculum seminis pear , oval

- 17 -

or bag shaped, right or left of the ovary. Shell glands diffuse.

Laurer's canal prominent. Uterus in coils from the ovary to the

ventral sucker. Vitellaria lateral, occupying the middle third

of the body and extending posteriad to the posterior testis. The

transverse vitelloduct passes from each region of the vitellaria

ovarial and unites before or agter reaching the ovary. The break

of the vitellaria in front of the ovary varies in extent from

0.06 to 1.9 mm. Eggs long oval in shape, cap distinct with an

operoular rim at the junction of the oap and the egg proper.

Length 0.0286 to 0.0357 mm by 0.014 to 0.0162 in width. The

excretory bladder is Y-shaped. The excretory canal sigmoid..

The genital pore at the anterior margin of the acetabulum. Skin

provided by minute needle like spinelets on the anterior portion

of the body as far caudad as the ovary. Spinelets deeply iu-

bedded in the cuticula.

Habitat - Biliary ducts of the domestic cat and a pet Coyote.

Distribution - Nebraska and Iowa - W. S. A.

Opisthorchis interruptus Braun (1901:5)

The animals are flattened, of band like form, the sides of

the body wavy, tapering anteriorly, posterior broader than the

anterior. The extreme posterior end pointed. 7 to 8 mm in length;

in the region of the acetabulum 0.5 mm in width; at the testes

0.7 mm . The oral sucker subterminal 0.25 to 0.26 mm broad,

acetabulum 1.35mm in diameter, and 1.3 mm posterior to the oral

sucker. A distinct prephajrynx is wanting, the pharynx is 0. 104

by 0.125 mm. The oesophagus has the same length as the pharynx.

The two wide intestinal crura extend to the posterior end, one

orus being longer than the other. The posterior testis lies in

- 18 -

front of the posterior body wall by a distance of 0.? to 0.8 mm;

diagonally to this the anterior testes; they are not af the same

size and form, the anterior four lobed, the posterior is larger

more nearly round and has less lobes. Between the testes runs

the S-shaped excretory bladder. The receptaculum seminis is

round or pear Bhaped and is smaller than the testes. Ovary median

of the same size as thetestes, oval in shape and lies just in

front of the receptaculum seminis. The uterus coils transversely

and is compact, the genital pore is situated immediately anterior

to the acetabulum. A ctorrus pouch is absent. The aggs are dark

colored, 0.023 by 0.01 mm in size. The small vitelline folliclef

have a grouping distinctly as in the other species of the genus

Opisthorchis; they begin anteriorly behind the acetabulum at a

different level on each side, posteriorly they reach to the

pesterior testis. The distance of the anterior end of the vit-

ellaria from the acetabulum is equal to the distance from the

acetabulum to the oral sucker. It is characteristic that in the

region of the receptaculum and as well in the ovarian region, an

apparently long break in the vitellaria appears on both sides.

This is approximately 0.6 mm long. The forms from the intestine

of Ardea viridirufa appear to belong to the same species as they

have the same length, the suckers and eggs are the same size and

in every other respeot correspond flbo 0. interruptus, the vitellar-

ia however could not be made out in those from Alcedo viresoens.

Habitat - Ardea viridirufa and Aloedo virescens.

Distribution - Brazil.

Opisthorchis lancea Diesing (1850:579)

The body is lanceolate in form, neck and trunk regions present

- 19 -

The length of the body 5.5 to 12.5 mm. The greatest width 1. to

2.8 mm. The neck region is one third the total length of the

entire body, is conical and elliptical in cross section. It

carries on its forward end a subterminal oral sucker. The walls

of the neok show a smooth wavy appearance, while that of the post-

erior end appears serrated. The oral sucker measures 0.33 to 0.36

mm by0.56 to 0.66 mm. Acetabulum in anterior third of the body

0.51 to 1.2 mm in diameter and is more spherical than the oral

sucker. A prepharynx is wanting. The oesophagus is short, in-

testinal crura reach to the posterior tapering end and run somewhat

parallel. The excretory system is Y-shaped, the main branoh wind-

ing in an S-shape between the testes to the excretory pore, which

is terminal. The anterior testis is situated on the left and has

four lobes, the posterior on the right with five lobes. The

cirrus pouch is absent. The genital, pore is situated median

immediately anterior to the acetabulum. The receptaculum seminis

is spindle formed, The ovary is biscuit shaped. The shell gland

is indistinct. The eggs are oval 0.029 to 0.033 mm by 0.012 to

0.014 mm. The vitellaria lateral to the intestinal crura, extend

from the posterior of the acetabulum to the fcevel ihers the crura

end. They are formed of small cross running acini, which com-

bine in more or less distinct groups. On each side there are

eight such groups of acini. The paired vitelline ducts begin

usually between groups 4 and 5 of the acini, a space being here

so that the vitellaria has two regions, and anterior ovarial

and a posterior ovarial region. Amphitypy is evident in this

species

.

Habitat - Male Delphinus tacuschi (Sieno tucuxi Gray)

Oroella breviroetria

- 80 -

Distribution - Barra do Rio Negro, Brazil.

Opiathorchia ovalia , Barker (1911:518)

General ahape of the body Blender and oblanceolate , the

anterior end tapering and bluntly conical, the poaterior end

broader and tapering alightly towarde the end, There is a well

defined narrower region or neck anterior to the acetabulum; the

length of the body meaaures 6 to 14.32 mm; the breadth aero a a the

acetabulum 0.343 to 0.81 mm; acroas the ovary 0.45 to 1.373 mm.

Small needle like, retroae spinels t a abundant over the anterior

two-thirde of the body, being moat abundant at the anterior end

as far caudad aa the acetabulum and becoming sparser until the

region of the ovary; in these speclmean these epineleta werequite

abundant. The oral aucker i8 8ubterminal 0.198 to 0.36 mm by

0.198 to 0.378 mm. The acetabulum in the anterior fifth, sixth,

or aeventh of the body, measuring 0.1331 to 0.216 mm by 0.131 to

0.325 mm. The genital pore immediately anterior and median to the

acetabulum. The pharynx 0.188 to 0.15 mm by 0.108 to 0.162 mm.

The oesophagua 0.09 to 0.63 mm in length. The inteatinal crura

are unequal in length and extend to the poaterior end. The tea-

tea are in the poaterior third of the body, oval ovate or

orbicular, amototh or 8lightly lobed. A cirrua ia abaent. Ovary

oval, ovate or orbicular, smooth or llightly crenate. The reoept-

aculum prominent, two to four times the size of the ovary.

Laurer'8 canal prominent. Shell gland diffuae and somewhat in-

die tinot; uterine coila of median density and compactnees.

Viteliaria lateral to the inteatinal caeca, divided into two

dee tine t region8 by a break near the ovary. Vitelline glands

begin 0.36 to 1.89 mm poaterior to the acetabulum and extend to

- 21 -

the posterior testis. Eggs short ovals in form, measuring 0.024

to 0.312 mm by 0.014 to 0.019 mm, with distinct lid but no oper-

cular rim.

Habitat - Bile ducts of soft shell turtles (Trionyx spinifer

Lesson and Trionyx muticus Lesaeur) and Chrysemys

matginata.

Distribution - Mississippi river in Minnesota and Iowa, U.

S. A.

Opisthorchis noverca Braun (1902)

Syn:- Distoma conjunotum Lewis and Cunningham 1872

Distoma conjunotum McConnell 1876

Distoma conjunotum Cobbold 1859

Body lancet shaped, spinous, 9.5 to 12.7 mm long by 2.5 mm

broad. Suckers close together, the oral being larger than the

ventral. Pharynx spherical, intestinal crura extending far post-

eriad. Genital pore immediately in front of the acetabulum.

Testes round or only slightly lobed, situated at the limit of the

posterior third of the body and nearly opposite. Ovary slightly

lobed, in front of the bifurcation of the Y-shaped excretory

reservoir. Uterus in loops through the central area from the

ovary to the genital pore. Vitellaria lateral, beginning at the

bevel of the acetabulum, extending to the middle of the posterior

testis. A break of the vitellaria occurs in front of the ovary

but it is indistinct. No cirrus sac. Eggs oval 0.034 by 0.021

mm •

Habitat - Gall duots of man.

Distribution - India.

- 22 -

Opiathorchia oaninua Barker (1911)

Syn:- Distoma conjunctum Lewis and Cunningham 1872

Opisthorchis noveroa Braun 1903

Parppiathorchia caninua Stephens 1912

Opiathorohis oaninua Barker 1911

Body aligntly lanceolate, length varie8 from 2.75 to 5.75 mm

in preeerved 8pecimena. Average length 3.6 to 5.2 mm. Body uni-

formly apinoae. No spines on the pediole. Oral eucker 0.28mm.

Pharynx 0.224 by 0.184mm. Oeaophagua 0.04 mm. The ventral

sucker 0.176 mm in diameter . n The acetabulum on the acetabular

papilla between the first and aecond anterior fifth; smaller than

the oral aucker. Pedicle about £ mm long. May be completely

retracted. Genital pore on the apex of the pedicle in front of

the acetabulum* Ita exact position variea with the atate of

contraction of the parts. In certain caaea it actually opens

externally to the aucker and anterior to it. The opening ia cov-

ered with acalea. The vas defarena and uterus run alongside one

another until they merge near the apex of the pedicle into a

common sinus. The testes lie in the poaterior third of the body;,

are orbicular or alightly lobed. One aituated obliquely in front

of the other. Ovary multilobular, the lobea 5 to 8, irregular in

size and shape. It ie located at the beginning of the poaterior

half. The aemiaal receptacle is globular, aituated to the right

of and doraal to the poaterior lobe of the ovary. Laurer'a oanal

U8ually runa from the end of the receptacle with a single curve

medially and backwards. The uterine coila form looaely packed

tranaverae coila terminating slightly in front of the level of

the first vitelline acinus. From here the uterua pas sea forward

- 23 -

into the pedicle to the left and ventral to the seminal vesicle.

Seminal vesicle commences about the level of the first vitelline

aoini. The coils displace the uterus ventrally and to the left.

In the pedicle the vesicle diminishes in extent and lies in its

dorsal (anterior) side .

Habitat - Liver of Pariah dogs.

Distribution - India.

Opisthorchis speciosus Stiles & Hassall 1894

Syn:- Distoma (Dicrocoelium) longissimum var. corainus

Stiles and Hassall 1894

Opisthorchis speciosus Stiles and Hassall 1896

Body elongated, its narrow and tapering anterior end covered

with retrose spinelets, length 7.6 to 29 mm by 0.666 to 1.333 mm

wide in the region of the ovary. Oral sucker subterminal , 0.291

to 0..366mm by 0.291 to 0.357 mm. Acetabulum in the anterior third

or fourthof the body, 0.178 to 0.244 mm by 0.178 to 0.244 mm.

The pharynx 0.15 to 0.205 mm by 0.131 to 0.188 mm. The oesophagus

0.15 to 0.30 mm long. Intestinal caeca equal or unequal, extend

to the posterior end. Testes in posterior eight of the body,

orbicular or oval, smooth or four lobed, Cirrus absent. Genital

pore not prominent, at the anterior mergin of the acetabulum.

Ovary in posterior fifth of the body, oval, ovate or elliptical.,

smooth to three lobed. Receptaculum seminis smaller than or equal

to the ovary. Shell gland diffuse or distinct. Laurer'e canal

prominent. Uterine coils very dense and close. Frequently ex-

tending laterally over the intestinal crura. Vitellaria divided

into two distinct regions by a break in front of the ovary.

Vitelline glands begin 0.003 to 0.263 mm posterior to the aoet-

- 34 -

abulum and extend to the posterior testis. Eggs long ovals, with

distinct lid and opercular rim, 0.085 to 0.033 by 0.015 to 0.018 mm

Habitat - Bile ducts of Crows (Corvus americanus, Corvus

ossifrage)and the frog.

Distribution - Maryland and District of Columbia, U. S. A.

Barker, F. D. (1911) divided the genus Opisthorchis and pro-

posed the generic name Amphimerus for those species which had two

vitellaria regions caused by a break in the line of acini near the

ovary, as postovarial and antovarial regions, while those forms

which had only one region he retained in the old genus Opisthorchis,

Genus Amphimerus Barker (1911:536)

Generic diagnosis - Fasciolidae; body flattened, elongated,

anterior end tapering. Skin frequently covered wholly or in part

by small retrose spinelets. Excretory system Y-shape with sigmoid

stem winding between the testes. Copulatory organs absent. Tes-

tes in posterior portion of the body, simple or lobate; the one

obliquely posterior to the other. Ovary anterior to the testes,

simple or lobate. Laurer's canal present; receptaculum seminis

well developed. Uterine coils anterior to the ovary and may

extend laterally over the intestinal caeca. Vitellaria well

developed, lateral to the intestinal crura and divided into two

distinct regions by a break op posite the ovary. Not extending

anteriorly befcond the acetabulum but frequently extending post-

eriorly to or beyond the posterior testis.

Habitat - Parasitio in the gall ducts of mammals, birds and

reptiles

.

Type species - Amphimerus oval is Barker (1911)

Stephens (1912) divided Opisthorchis once more retaining the

- 25 -

generic name for all the species exoept Opisthorchia oaninus

Barker which he placed under a new genus Parapisthorchis

.

Genus Paropisthorchis Stephens (1912)

Structure as in Opisthorchis; exoept that the ventral sucker

and genital pore oocur on the ape* of a precesa or pedicle pro-

jecting from the anterior portion of the body. This process is

about 0.5 ram long and is retractile.

- 36 -

?. DISCUSSION

" It is a fact well acknowledged by persons who have worked

with species of the genus Opisthorchis, that the forms in this

genus represent one of the most difficult problems to the

systematise, writes Looss (1899). * Several forms anatomically

seem identical that should be classified as separate individuals

on a biological basis" (1907). Opisthorchis felineus and O.gem-

inus without knowledge of their host will be classified under the

same species and in fact I am inclined to think 0. tenuioollis is

at most a variety of 0. felineus. 0. felineus is found in the liv-

er of man and the oat in Europe; but 0. geminus in Egypt in the

liver of the non-migrating Milvus parasiticus, Anas boschas fera,

Circus aeruginosus. 0. tenuioollis from seals in Europe. Neat

to these three forms comes O.viverrini and O.entxi which differ

from the above not in large details. Looss (1899) takes 0.

geminus as a separate speciAs on biological grounds. The above

mentioned birds are native to Egypt and never migrate and if they

are the same as 0. felineus, cats and man should be harboring

these parasites, however this has not been reported not has

Looss been able to find any Opisthorchis in mammals contrary to

his special search. 0. geminus differs from the rest on the

same ground as it differs from 0. felineus. Looss again makes a

comparison between 0. tenuioollis and 0. geminus from Anas boschas

fera. 0. geminus fron Anas is 10. to 12.5 mm long and 1. to 1 .7

mm broad; testes both lobed in 0. tenuioollis and 0. geminus.

Ovary of 0. geminus from two different kinds of hosts varies

slightly from that of Anas, being round or slightly lobed, but

- 27 -

from Circus more distinctly lobed. Vitellaria same both in 0.

geminu8 and 0. tenuicollis. The eggs of 0. temiiccllis as given

b y Mtthling 0.035 to 0.028 mm long and 0.013 mm wide, in gem-

inus 0.037 by 0.013 ram in Anas and 0.023 by 0.013 mm in Circus.

By such comparison Looss finds these forms identical yet thinks

they cannot be classified as one species for the same reason as

0. felinmis cannot be the same as 0. gem inus , because of the spec-

ificity of the host. The same argument is brought about in regara

to the nearly identical forms of 0. tenuicollis and 0. felineus.

Braun in comparing the forms found by Mtlhling with 0. felineus

comes to the conclusion that they are different forms.

0. entzi has very identical characters with 0* tenuicollis

which in turn is c&oeely related to 0. felineus and 0. geminus.

It is taken as a separate species however by only specific

differences of the euckers, in 0. entze the acetabulum ts larger

than the oral sucker which in other forms is equal to it or the

oral sucker is larger than the acetabulum. 0. viverrini differs

from the above mentioned form by the distinctly ifcbed ovary, by

the moderately developed uterine coils and by the egg with an op-

ercular rim. No figure aocompanies Von Ratz's description of 0.

entzi whioh makes a comparison with 0. viverrini less valuable.

The forms discussed above differ from each other by one or

two characters in general. Two other forms which are more

distinctly separated from the rest are 0. longissimus and 0,

etaulans. Both of these have ovary very much lobed (follicular) .

In the figure of Looss (1891:52) this character is not noticeable

but his figure in a later description (1899 :673) gives these

characteristic lobes. The difference between O.simulans and 0.

- 38 -

longissimus is quite small. 0. longissimus tapers anteriorly tut

0. eimulans only slightly. The reverse is the case posteriorly.

The oral sucker in the later is much larger than the acetabulum,

the reverse is the case with 0. longissimus. The uterus coils

are much closer in 0. longissimus than in 0. simulans. Testes in

the former larger and somewhat lobed. Located more posteriorly

than that in 0. eimulans which is more round, smaller and not

not so much posterior, some of these characters are much in var-

iance as I have described in the case of 0. viverrini var.

Chiengmai. Yet the variations in the same species are with in

normal limit.

Opisthorchis obsequens Nicoll is distinguished by the smaller

form, by its very much lobed testes which suggest the genus

Clonorchis. The ovary is usually irregular in outline, ventral

sucker is larger lihan the oral sucker; vitellaria more continued.

0. piscicola can be readily distinguished from the rest of

the species by its minute size, with its relatively larger

acetabulum. In this respect being much like 0. obsequens but

differs from it by its broad spines, lobed testes and its piscine

host

.

Another form described by Galli-Valerio (1898:145) and

named as pianae is questioned by Kowalewski (1898: 751).

Certain characters were overlooked which will make these forms

identical with E. conoideum.

Galli-Valerio (1898:933) tho admitting that this form

may not belong to the genus Opisthorchis maintains that it is

not identical with El conoideum. From the description he re-

gards it as Opisthorchis as it agrees with 0. Blanchard.

- 29

-

From his figure it is certain that it does not belong to the

genus Opisthorchis as the vitellaria extend to the posterior end

of the body in an H-shape. However I cannot pass judgment re-

garding this form being an Amphimerus from his meager description.

A 8 Kowalewski has made a careful study of these forms it is safe

not to include 0. pianae in even Amphimerus.

Kowalewski (1898:52) described a form 0. simulans var.

poturzycensie from the liver of Anas boschas dom. He makes a

comparison of this with 0. simulans Looss and 0. longiesimus.

His comparison is based upon the description of Looss (1896:52).

Loose (1899:674) describes another form of 0. simulans with slight

variation from Anas penelope and Felix crista ta. This form slight-

ly larger than that from Pernis apivorus corresponds to Kowalew-

ski* s drawing (Fig. 23) and description.

Barker's division of Opisthorchis into two genera is question-

ed by some parasitologists. Leiper( have not been able myself

to consult Leiper^ attiole on this subject) has objected to such

division on the ground that extent of the vitellaria posteriad -

is a matter of chance and not a generic characteristic. I have

examined more than fifty speoimens of 0. viverrini and I find the

variation in the extent of the vitellaria small. They never ex-

tend behind the anterior testis, usually the vitellaria end at the

level of the ovary. I have never noticed any break in the line

in that region. The number of acini, their anterior or poster-

ior extent, and the number of acini behind the transverse vitell-

ine duct, does vary somewhat but always within a normal limit.

Muhling (1896:261) examined sixty specimens of 0. felineus

and from the description I find the variation in 0. felineus the

* 30 -

same ae in 0. viverrini. 0. felineus is the type species of the

genus Opisthorchis . I find from the descriptions the same thiBg

true for the vitellaria in all species of thia genus as modified

by Barker. The other species of Opisthorchis are classified und-

er the genus Amphimerus Barker. This new genus is based entirely

upon the division of the vitellaria into two regions, an antovar-

ial and a postovarial with a distinct break in the line of acini

at the ovary

.

Weski (1900) examined 400 specimens of 0. lanoea Dies, for

the extent of the variation of the vitellaria. In every case

the vitellaria did extend caudad to the posterior testis which

never was the case with 0. felineus or the like. There was some

irregularity in the line of acini opposite the ovary. In twenty-

two cases only there was no break in the line of acini opposite

the ovary on the right and only four on the left, but the acini

were distinct. In 119 cases the groups coalesced on the right

and in 35 on the left.

Ward (1894) describes the vitellaria in Opisthorchis pseudo-

felineus as follows, "Vitellaria extend to the middle of the

space between the two or even as faaj as the anterior edge of the

posterior testis. One break in the line of acini may always by

recognized as most prominent; it is located just opposite the ovary

on each side, and is in length more or less equal to the diamete-r

of the ovary. This is the condition shown on the left side of Ithe

worm figured. This space divides the vitellaria into two portions

which may be distinguished as antovarial and postovarial. While

it is in some cases possible to distinguish in the antovarial

portion groups of acini they seem to be usuallyrather indistinct

- 31 -

or at least very unequal in size, as if adjacent groups had become

confluent "by the growth of interlying acini. The posterior por-

tion, however is usually distinctly divided intctwo or three

groups of acini, though even these may he obliterated. In two

or three oases a small group of acini was found, on one side only

in this intermediate space opposite the ovary, and was clearly

separated from both antovarial and postovarial portions by a

small space. This was the case in the right vitelline gland of

the worm figured, corresponding to the two portions of glands,

one finds on each side two ducts which, extending obliquely to-

ward the ovary from a short distance before and behind it, form

a "Y" or "V" according as they meet before or are not united

after reaching the ovary. These ducts on dither side of the ovary

form one of the most characteristic appearances of the specimen".

Stiles (1904:32) describes these glands, w the antovarial

portion four acini, postovarial three, each portion provided

with its own oviduct, the antovarial portion extends from the

ovary oephalad to about the boundary between the anterior and

equatoral thirds of the body; the posterior portion extends

caudad from the ovary to about the anterior plane of the post-

erior testis."

Barker (1907:100) makes a detail description of the var-

iation of the vitellaria in 0. pseudofelineus from more than a

hundred specimens.

"The vitellaria never extend to the acetabulum, but their

position posterior to posterior matgin of the acetabulum varies

on the right sied from 0.3mm to 4.5 mm, the mode being 1.1 mm

with frequency of 23 per cent. On the left side the position

- 32 -

varies from 0.3 mm to 3.7 mm the mode being 0.7 mm with a fre-

quency cf 18 per cent*****************************************

*********** The extent of the vitellaria caudad varies from the

anterior margin of the anterior testis to a place 0.07 mm to

0.30 mm posterior to posterior testis, with a frequency of 28

per oent for the right side. On the left side the extremes

were found to be a plane 0.16 mm anterior to the posterior tes-

tis the mode being the anterior margin of the posterior testis,

with a frequency of 22 per cent."

From the above quotations we find that the variation in the

extent of tie vitellaria in 0. pseudofelineus is great but the

break in the line of acini at the avery quite constant. With all

the variations 0. pseudofelineus represents a homogeneous type

which stands separately. Forms belo»ging to this type are 0. inter

ruptus, 0. lancea, 0. noverca and 0. specie sue and 0. oval is, all

having vitellaria divided into regions and a distinct break tak-

ing place at the ovarial region. In 0. noverca the same is true

except that the break in the line of acini at the ovary is less

distinct. In this later form the vitellaria extends from lateral

areas behind the acetabulum to the posterior testis.

Barker (1911) mentions that Amphimerus noverca is the only

form which does not present a break in the line of acini in froat

of the ovary. But rightly he questions the accuracy of McConnell's

drawing. It seems that Barker has not noticed the figure given

by Leuckart. In this there is a distinct break in the line of

acini in front of the ovary on one side.

The topographic details of the two groups are identical, they

are parasitic in the same animals in the same organs. A study of

- 33 -

vitellaria among distomes in general shows forms in which the

vitellaria varies from a knob like gland on each side to forms in

which the vitellaria surrounds the whole body under the integument

except a median dorso-ventral apace. A typical type seems to b»

H-formed as is seen in 0. lancea. It seems variation in the ex-

tent of the vitellaria comes by the degeneration of the glands

as one of the possible causes. I have noticed in few specimens

of 0. viverrini, Clonorchis endtmicus and Clonorchis sinensis that

one or more acini are missing but the longitudinal duct is stand-

ing full of shell material. In other forms there were one or more

acini below the transverse duct on one side and in others on the

other side. The general position of the transverse duct was con-

stant, though the ducts coming from each group of acini vary con-

siderably. In 0. pseudofel ineus the vitellaria has four possible

combinations both sides "V" shaped or "Y" shaped, right "

V

w, left

WY" shaped or the reverse. The position of khie transverse duct

being constant, I find it is in the same place in both groups.

Having all these variations and objections for the formation of a

new genus Amphimerus, I find more argument in favor of such a

division than against it. The genus Metorchis is based chiefly

on the extent of the vitellaria anterior to the acetabulum, the

genue Clonorchis entirely on the dendritic testes, it is true

that vitellaria e«tent posteriad varies more than anteriad but ev-

en so there is great constancy in this posterior extention. Among

forms which form a homogeneous group 0. obsequens Nicoll repre-

sents as much an intermediate form to the genus Clonorchis as the

variation in the extent of the vitellaria does in Opisthorchis

.

Thus the forms 0. pseudofel ineus , 0. interruptus, 0. lancea, 0.

- 34 -

noveroa, 0. ovalie, and 0. spec io sue became Amphimerus pseudo-

felineus, Amphimerus interruptus, etc,.

Amphimerus pseudofelineus , A. oval is and A. speciceus are the

three American species very much alike. Barker (1911:520) makes

a detailed comparison based chiefly upon measurements some of

them to tbBtks of a micron* Some of these measurements are con**

fusing since a great range of variation is represented. The oral

sucker in A. ovalis varies from 0.198 to 0.36 in length and in

width from 0.198 to 0.378 mm, egg from 0.024 to 0.812 mm long by

0.014 to 0.019 mm wide. The mode of preserve tion might be one of

the causes of variation in measurements. I find the egg one of

the most constant characters in 0. viverrini, 0. felineus and 0.

pseudo-felineus , there are some abnormal eggs which must be

rejected in measuring. The topography of these three American fo

forms seem very much alike. Especially A. ovalis which is des-

cribed as a new species chiefly because of the specificity of its

host and some minor differences. Recently I have had for study

a single specimen collected from Chrysemys marginata Agassiz

taken near Fairport Iowa, which although poorly preserved, cor-

responds to the description of Barker except that the uterine

coils are more compact, as in A. spediosus. Some of these dif-

ferences according to Barker go more in variance among the indiv-

iduals of the same species than between the species themselfes.

A. ovalis differs from the rest in its smaller size, the shape of

the body being more slender, the oral sucker smaller, oesophagus

larger, uterine coils more dense. Amphimerus specicsue has

more characters in common with A. pseudofelineus than with A.

ovalis; it differs however by the small size of the oral sucker,

- 35 -

the position of the ovary (at the posterior fifth of the body) and

its large size; by the more densfcy coiled uterus and by its avian

host. A.ovalis in general is smaller than A. pseudofelineus and

A. speciosus ia more slender has smaller oral sucker, larger

oesophagus, position of the ovary more anterior (posterior half

of the body), less dense uterus coils and smaller egg, constant

oval shape of the ovary and testes and the reptilian host.

Amphimerus interruptus differs from the three American forms

by its small size, 7 to 8 mm, its lobed testes, smaller and elong-

egg, notably the difference in size of the oral and ventral suckers

the former being much larger.

A. noverca once so much confused ia easily distinguished

from the above closely related forms by its general form of body

which is smaller and broader, its very pocrly developed uterine

coils, its broad spinelets and the vitelline glands which are

indistinct

.

MoConnell (1876:342) and (1878:476) found dietomes in the

biliary ducts of a Mohammedan, twenty eight years old who died

in the medical college of Calcutta; and again in 1878 from another

Mohammedan twenty four years old. McConnell diagnosed these

forms as D. conjunctum Cobbold (1858). Lewis and Cunningham

(1872:168) found in Pariah dogs in Caloutta, forms which they

diagnosed as Distomura conjunctum. Stiles and Hassall (1894:430)

showed that McConnell • a form differed from that of Lewis and

Cunningham. Looss (1899:565) placed Cobbold 1 s Distomum conjunctum

from the American red fos, Canis vulpis in the genus Metordhis.

Eraun (1903: 164) gave to McConnell 'a form the name 0. aoverca.

Barker (1911:532) placed that of Lewis and Cunningham in the

- 36 -

genua Opisthorchis and named it 0. caninua . Stephens (1912)

placed 0. caninus in a separate genus; Parepiethorchis . This new

genus is based upon the 0.5 mm pedicle upon which is situated the

acetabulum and the genital pore. In all forms of the genus Opisth-

orchis, the acetabulum is a cup shaped sdsaile adhesive organ

sunken in the body parendhyma. In Paropiskhorchis hawever the

acetabulum and the genital pore are on a pedicle which is re-

tractile, such a character is a generie one. In all other respect

the genus Paropisthorchis is like that of Opisthorchis.

Trom the above discussion one thing is clear; that is the

need of experimental woBk to clear the specificity of many of

these forms. I am inclined to think such a work will bring the

identity of 0. geminus, 0. felineus, 0. tenuicollis and 0. entzi

under one species. A. ovalis and A.speoiosus in another. Such

an experiment can be done as suggested by Looes (189S) by feeding

experimental animals. Another thing that will do much in proving

or disproving the identity of these forms is careful search among

hosts in question and a study of the life cycle in relation to

the mode of transmission.

Key to the genera of the subfamily Opisthorohiinae

Loose 1899.(See page 2 for diagnosis of the subfamily)

1. (2) Vitellaria lateral to intestinal caeca - - — 3

2 . $5Vitellaria anterior to intestinal caeca, lateral to

oesophagus uterine coils fill entire body post-

erior to acetabulum HOLOMETPA

3. $2) Vitellaria not Mtending anteriorly beyond the

acetabulum --------- - — ------4

- 37 -

4 . Vitellaria extending anteriorly beyond the acetabulum

METORCHIS

5. (6) Vitellaria not divided into two distinct regions

by a break in the line of acini near the ovary - 7

6. (5) Vitellaria divided into two distinct regions by a

b break in the line of acini near the ovary -

7. (8)

8. (7)

C10N0RCHIS

9* (10) Acetabulum and genital pore on a pedicle

PAROPISTHORCHIS

10 . 0) Acetabulum and genital pore not on a pedicle-

0PX8TH0RCHIS

Key to the species of the genus Amphimerus

Barker 1911

1

.

(3)

2

.

(1) Testes generally simple, oval - - A. ovalis

3

.

Coils of the uterus moderately developed - 5

4 . (3) Coils of the uterus well developed A. 3peciosus

*. (6) Number of acini in postovarial portion of vitellaria

6. (5) Number of aoini in postovarial portion of vitellaria

7

.

(8) A break in the line of acini moderate A. pseudofelineus

8. (7) Break in the line of acini indefiniteA. noverca

9. (10) Postovarial acini very marked - A. interruptus

- 38 -

1©. (9) Braak in the line of acini moderate A. landea

Key to the speciee of the genua Opisthorchis

1. (2) Body spineless - - - - -

2. (l) Body covered with spines 0* piscioola

3

3. (ft) Suckers unequal, uterus well developed - 5

4. (3) Suckers equal uterus poorly developed

0. viverrini

7. Oral sucker larger than acetabulum 0. simulans

8. Oral sucker smaller than acetabulum

0. longissimus

9. (5) (12) Oral sucker smaller than the acetabulum- -10

10. (11) Found in the liver of Mammals - 0. tenuicollis

11. (10) Found in the liver of birds - 0. entz*

12. (5) (9) Oral sucker larger than the acetabulum - 13

13. (14) Found in the liver of Mammals - -

0. fslineus

14 (13) Found in the liver of Birds - 0. geminus.

5. (6) Ovary not multiiobed

6. (5) Ovary multiiobed - -

9

7

- 39 -

V. DESCRIPTION OF OPISTORCHIS 7IVERRINI

POIRIER 1886 VAR. CHIENGMAI

The general diagnosie - This parasite is leaf shaped, ob-

lanceolate flattened dorso-ventrally . The preserved specimens

are almost transparent, uterus region somewhat dark and always

full of the numerous yellowish tinged eggs; the body has many

ridges mostly passing transversly on account of the contraction.

The integument is roughened by rubbing. The anterior third of

the body starting at the acetabulum gradually tapers and is al-

most conical at the anterior end. At the le?el of the acetabulum

on both sides of the body there is a slight shallow constriction

which distinguishes a neck region. Posteriot to the acet abulum

the lateral borders run fairly parallel until to the posterior

testis, from there it gradually decreases in width till at the

extreme posterior end it tapers bluntly. The length and breadth

vary according to the contraction and normal individual variation.

The length varies from 3.7 to 7.7 mm. Two specimens are 6 mm in

length, two 5, three 5.4 mm, four 4.6 mm. The average length of

twenty specimens is 5.3 mm.

Width and thickness varies from 1 to 1.3 and 0.83 to 0.44 mm

respectively. The suckers are about 3/7 of the length of the

body distant from each other. The variation is largely due to

the expansion of the cephalic tegion at time of fixation. The

suckers are about equal size 19 by 23,4 mm. The length of the

oral sucker varies considerably from 0.125 to 0.23 mm. The Width

from 0.17 to 0.26 mm and 0.19 to 0.27 mm respectively. The acet-

abulum measures 22.5 by 23 mm . The oeslphagus is much longer

- 40 -

than the pharynx, the former 0.31 mm on the average varying from

0.24 to 0.42 mm, the latter 0.14 by 0.13mm. The intestinal crura

reach almost to the posterior border and are often filled with

bWdd. The excretory pore is at the posterior extremity and the

excretory bladder curves between the testes in an S-shape. The

testes are located at the posterior, approximately 2/7 of the

bGdy length from that end. They lie obliquely one bshind the

other; the anterior one has four lobes, the posterior one five.

The ovary is multilobed with smooth margin; it occupies the ant-

erior part of the posterior J of the body (average) slightly to-

ward the right (0.08) from the median. Behind the ovary lies

the pear-shaped or retort-ahaped seminal receptacule. Laurer's

canal present. The uterus coils occupy a median position in

the middle third of the body. The vitellaria occupy the faitly

broad lateral areas, in about the middle five, six and seven

and a half tenth's of the body. The number of acini varies from

7 to 9 usually 8 on each aide. It never passes posteriorly be-

yond the anterior margin of the first testis nor reaches to the

acetabulum. Cirrus pouch absent. The eggs are oblique oval with

a round operculum at the sharper end 0.026 by 0.0156 mm. There

is a distinct narrowing towards the anterior end, a distinct

lid followed by a shapply projecting brim as in Clonorchis sinen-

sis .

Habitat - Liver.

Host - Indian civet oat, first tim» reoorded from man in

Siam 1915

I included the above short description in order to give a

general idea of the form; but it seems to me tkat minor details

- 41 -

need to be mentioned though some of them are in considerable

variance, for easy diagnosis by later workers with the same kind

of forms. All present confusions would have been eliminated if

farmer investigators had given a detailed description of this form

The oral opening of 0. viverrini var. Chiengmai is nearly

circular in transverse section, terminal and surrounded by the

anterior sucker. The thiokness of the oral sucker is about 0.13nm.

Behind this there is a small poeket immediately followed by a

circular pharynx 0.41 by 0.13 mm. The length varying from 0.13

to 0.15 mm, width constant. The length of the oesophagus in some

cases is three times as much as the pharynx but this region is

much at mariance; from 0.24 to 0.42 mm with an average of 0.31 mm.

At the base of the oesophagus there is an enlaggment before the

bifurcation takeB place. The intestinal crura extends between

the uterine coils and vitellaria nearly to the posterior end of

the body. Usually both end at the same level. Sometimes the light

or left extending further. The average distance from the posteiAor

end to the end of the crura is 0.25 mm, varying from 0.11 to 0.4

mm •

The female reproductive organs are composed of multilobed

ovary, the vitellaria on both sides of the margin with a trans-

verse duct. The shell glands are composed of isolated cells ope»

ing into the ootype. The uterine coils dorso-ventrally and right

to left in front of the ovary, and opens into the common genital

pore immediately anterior to the acetabulum.

The vitellaria occupies a median postion of the body from

0.03 to 0.13 mm from the body margin on both sides . fhe number

of acini is regularly 8 on each side though it varieB from 7 to 9

the extra one usually being on the right side. The right vitellaia

is larger than the left, it starts 0.38 mm from the posterior end

of the acetabulum on the average. The closest it extends ant-

eriorly to the posterior end of the acetabulum is 0.24mm, the

fartherest 0.9 mm. The left vitellaria 0.43 mm behind the acet-

abulum on the average varying from 0.3 to 0.84 mm. The range of

extent of the vitellaria posteriorly varies little. It never

extends behind the anterior testis more than 0.4 to 1mm on the

right, 0.43 to 0.06 6n the left. Taking the ovary for the

comparison of vitellaria extent it is 0.06 to 0.42 mm below the

anterior end of the ovary averaging 0.2 mm. Each acinus is con-

nected by curved longitudinal ducts on each side. Later this

duct unites with the transverse <Juot forming a broad V-shaped

figure. The upper duct unites with a duct coming from the lower

acini in front of the vitelline glands the united duct farrms the

transverse duct. The same kind of union takes place from the

left vitellaria but not always in regular manner as sometimes the

union from the upper and &ower ducts takes place between or afte r

they leave the glands.

The shell glands are somewhat diffused on both sides of the

ovary and in front where the uterus begins. They are single iso-

lated cells scatered in the parenchyma. They open into the ootype

which is a little above the region where the vitelline duct opens

into the uterus.

The ovary is multilobed, its margin smooth slightly toward

the right (0.44 mm from the right side of the body 0.52 mm from

the left). It occupies the anterior part of the posterior one-

fourth of the body. The average width of the ovary is 0.46 and

the length 0.22 mm varying from 0.34 to 0.57 mm by 0.19 to 0.29 mm

- 43 -

respectively . The number of distinct overy lobea varies consid-

erably from 3 to 9. Some of the lobes are much larger than others.

There is alwahs a regularity in having two branches, a right and

a left one on eaoh side of the receptaculum seminis resembling a

saddle. The ovary is more antero-ventral to the receptaculum

seminis and the lobes have three directions, right, left and med-

ian. In 80 per cent of the specimens examined the right and left

lobes are equal. The oviduct opens into the uterus below the vit-

elline duct. Just above the common branch of Laurer*s canal and

the receptaculum seminis. The uterine coils occupe a median

position in the middle 1/3 of the body. The uterus is filled

with nature yellowish eggs. The lower end unites with the

ootype and is mostly filled by spermatozoa. The genital pore

is immediately anterior and median to the acetabulum, it is some-

times slightly toward right or left. The uterus opens in the

common genital atrium by the side of the vas deferens.

The vas deferens ooils from the right side and opens into tbe

common atrium at the side of the female genital pore. The vasa

dfferentia coming from eaoh testis unite between the uterine

coils and form the vas deferens. The two testes are placed ob-

liquely one behind the other. The anterior testis is fonn lobed,

the posterior five lobed. The lobes are not always regular.

From twenty specimens only one anterior testis had three lobes,

in three the lobes were poorly developed the rest were well

developed. There is greater irregularity of the posteriM testis

than the anterior one. In three specimens there was only four

lobes, in another three, four others had five fcobes but one hr

two of these were very poorly developed. The anterior testis

- 44 -

is more to the left aide of the body, the posterior more median

and somewhat inolined toward the right. Both testes are elevated

from the lefel of the body and occupy most of the dorso-ventral

apace. The anterior testis is located in the 6th or 7th posterior

tenth of the body at the posterior 3/7 of the body. Its length

varies from 0.19 to 0.48 mm. The average 0.34 mm. The width

from 0.22 to 0.48 mm. The average 0.33 mm, the length and width

of the posterior testis is 0.36 by 0.47mm, varying from 0.19 to

0.48 mm in length and 0.27 to 0.48 mm in width. There ie a marked

regularity in the position of the lobes, the anterior testis has

four lobes and in 80 per cent of the cases these lobes are ar-

ranged right, left anterior and posterior. The posterior testis

has five lobes and theae are arranged anterior right, anterior

left, right, left and posterior in 60 percent of the cases. In

20 per cent of the cases for anterior testis the tendency was

anterior right, anterior left, posterior right and posterior

left. In 30 per cent of posterior testis the tendency was anter-

ior, anterior right, anterior left, posterior right, posterior

left, in the rest of 10 per cent irregular.

The seminal receptacule ia of medium size, pear or retort-

shape and dorsal to the ovary; in all cases filled with sperma-

tozoa. Laurer's canal starts from the common duct into which

the receptaculum seminis opens and extends downward, curves

backward, then to the right making a larger diameter to the tube,

then it nuns straight downward and opens on the dorsal surface of

the body on the dorao-lateral aied of the receptaculum seminis.

The excretory bladder is located in the posterior fourth of

the body, it statts a little behind and lateral to the reoept-

- 45 -

aculum seminis and latero-poeterior to the ovary. It curves

between the testes in an S-shape then enlarges greatly in diameter

forming a buld behind the posterior testis, it then narrows con-

cise rably and opens to the outside thru a narrow slit. The dia-

meter at its anterior portion is about O.48 mm, between the testes

0.23 mm, at the bulb region 0.4 mm at the narrow nedk 0.15 mm,

and at the pore 0.01 mm. These figures vary considerably.

The reproductive system - The genital pore opens immediately

anterior to the acetabulum 0.008 to 0.018 mm. Mostly median though

in some oases slightly to the tight or left. The genital cloaca

is about 0.07 mm long. The cloaca is flask shaped. The metra-

term, the terminal portion of the uterus, has the same diameter

as the cloaca at this region which is a continuation of the first .

The ductus ejaculatorue narrows into a vary narrow opening about

one-half to one-third the size of the metraterm opening and at

the side of it. The genital cloace gradually narrows like the nock

of atottle. The genital pore opens at the level of the anterior

of the acetabulum on the sentral side which is raised from the

general level of the body. The wall of the cloaca is surrounded

by heave muscular annular muscles . Numerous prostate glands are

located around the genital pore and open into it.

The vas deferens is filled with spermatozoa. It curves

with the uterus side by side dorsally and opens in the genital

cloaca 0. 06 mm from the ventral surface or fro$ the genital pore.

The vas ##ferens starts from the anterior testis on the inner sur-

face of the anterior lobe about 0.05 mm from the dorsal surface,

it curves slightly above the testes and curves in at the shell

gland passing between Laurer's canal and the intestinal crura and

- 46 -

under the transverse vitelline duct. The other vas efferens

starts from the inner anterior right lobe of the posterior testis

and joins the first at about 0.38 mm from the genital pore form-

ing the vas deferens. The diameter of the vasa efferentia is

about 0.01 mm the diameter of the vas deferens varies from 0.06 to

0.12 mm. The vasa efferentia unite between the uterine coils

slightly (0.09 mm) dorsal to the median portion of the body.

The vas deferens curves Centrally at the anterior margin of the

acetabulum toward the dorsal side of the acetabulum . At the

postero-dorsal region of the acetabulum it coils up till the post-

erior 1/3 of the acetabulum. Then it curves to the right on the

sane level and makes seven dorso-ventral loops before it reaches

the vas efferens. The loops are situated onthe right on the

same level from the median longitudinal plane of the acetabulum.

The loops ars found in two regions ane to the extreme right and the

Other more 4c the posterior. The extreme right loops are proximal

to the acetabulum, they coil dorso-ventrally much more and are

larger in diameter. The coils comeing from the posterior of

the acetabulum bend toward the right at about the same longitud-

inal plane with the lateral right margin of the acetabulum then

coila dorsally. From there a short continuation to the posterior

and to the right at that point it ourves to the ventral surface

then makes a short transverse landing. The last two coils are

short and at the same level as the provious one. The posterior

part of the vas deferens is straight and at the same level as

the extreme left extention of the anterior coils.

The testes are obliquely placed on the posterior 2/7 of the

body (average) . The anterior testis more on the left and the

- 47 -

posterior more median or slightly toward the right. The lobes

are short with round smooth margins. They originate from a commnn

center. The length of the testes'lobee vary from 0.095 to 0.15 mm

The median portion of the testes filling nearly the dtorso-ventral

space at this region and are slightly closer to the dorsal than

to the ventral surface.

The female reproductive system - The vitellaria occupies the

lateral borders between the margins of the body and the intestinal

canra. The vitellaria never extend cephalad to the acetabulum

nor oaudad. Only in two cases do the vitellaria extend to the

anterior margin of the anterior testis. The vitellaria do not

quite occupy the middle 1/3 of the body margin. The mean relation

in twenty speoimens examined gave 0.42 mm anterior non-vitellaria

portion, 0.34 mm median non vitellaria portion and 0.24bnm post-

erior non vitellaria portion. This ratio is in gonaiderable

variance in some individuals as the anterior portion extends some

what in fixation.

The vitelline glands are oval or spharioal, transversly

placed in the parenchyma. A longitudinal duct from both vitell-

aria having ducts opeaing from each gland and later th» two long-

itudinal ducts joined by a transverse vitelline duct which bends

obliquely to the ovary and slightly toward the left of it enlarges

forming the vitelline reservoir. In eefenteen out of twenty

specimens, the right vitellaria has 8 acini, 9 specimens show

only 7, 16 have 8 acini on the left, two have 9 and another

pair have 7 acini. At the beginning of the transverse duct, the

longitudinal duct ooming from the anterior and the one coming

form the posterior form a long Y-shaped fttgure. On the right in

- 48 -

suven forms the union is among the glands, in the rest 0.038 to

0.057 mm. toward the ovary. On the left in 9 forms the union of

the ducts is at the glands, the rest from 0.006 to 0.057 mm

toward the ovary. In six forms the rifcht acini were entirely

distinct; on the left in one form siven acini were in combination,

in two 5 acini and in three only two acini were in combination.

In three specimes only three acini are in combiaation on the

right. The space between the acini varies considerably but at

a small margin from 0)05 to 0.1£ mm the average being 0.09 mm.

In six forms the number of acini posterior to the transverse

duct on the right was tko', the rest, one in the same region. On

the left eight specimens have two acini the remainder having one.

There is no break between anterior and posterior pjar_t

.

The course of tie longitudinal vitelline duct is at the inner

margin of the glands median to dorso-ventral surface. Between

the spaces of acini it curves toward the intestinal orura, the

concave side being toward the glands. The width of the acini

varies from 0.09 to 0.14 mm the length from 0.18 to 0.35 mm. In

one case only an entire acinus was degenerated.

The transverse duct passes dorsally to intestinal orura.

The transverse duct from both sides unites between the ovary and

the seminal receptacle slightly toward the right. Branched

efferent ducts from these groups of cells unite with the longi-

tudinal ducts and these later with the transverse.

The vitellaria consists of numerous oblique oval, elliptical

lobes with smooth rounded margins. In some species the lobes

are very closely aggregated in others more or less distinct.

Each lobe is surrounded by a thin membrane. The lobes are held

- 49 -

in the parenchyma. They contain numerous cells of shell forming

material. These cells are mostly spherical and contain numerous

yellow granules. There is a vesicular nucleus well stained with

Erlich'e acid hematoxylin. It has a distinct nucleolus and a

number of chromatin granules.

The fitelline reservoiB is on the left side of the ovary

below the shell glands. The unpaired duct originates from the

ventral anterior side of the vitelline reservoir. At the proximal

end it proceeds anteriorly having a diameter of 0.018 mm at the

distal and, where it opens into the common duct 0.04 mm in diameter

The unpaired duct opens behind the ootype, at this point it has

a diameter of 0.06 mm. This measurement varies according to

the amount of yolk found in the duots. The thickness of the

vitelline reservoir is 0.045 mm. It is formed by the expansion

of the transverse duct in this region. The ieanswerse duct

continues dorsally while the unpaired duct below this proceeds

at right angles to the transverse duct and anteriorly toward the

ootype. This unpaired .duot is in a plane 0.02 mm ventral to that

of the transverse duct. The walls of the duct are homogeneous,

no nucleus is seen. The thickness of the duct walls is 0.003 mm.

The ovary is multilobed and has a smooth margin. It

occupies an anterior lateral and slightly antero-ventral position

to the receptaculum seminis. The average length is P2 mm and the

width 0.45 mm. The thickness varies according to the different

lobes. The antero-ventral lobes are thickest 0.2 mm the others

0.12 mm. The oviduct proceeds anteriorly 0.01 mm to the right

from the starting point where it jions the common duot behind the

unpaired vitelline duct. The length of the oviduct is 0.36 mm.

- 56 -

It is 0.072 mm ventral to the receptaculum seminis. The duct

coming from the receptaculum seminis and Laurer's canal curves

making a small arch above the Receptaculum seminie, then forma a

trough and begins to run straight anteriorly joining the oviduct.

The oo type is at the posterior end of the uterus and is a slight

enlargement in the diameter of the uterus. IK the ootype the

shell glands pour therr yolk and here fertilization of the egg

takes place by the spermatozoa which pass from the receptaculum

seminis uterinum into this region. The ootype is 0.12 mm fcom

the ventral surface and 0.23 from the dorsal, its thickness 0.013

mm on the average. Its distance from the receptaculum seminis is

0.072 mm. The walls have thich annular muscles. They have nuc-

leated cells. The lower part of the ootype is covered on the in-

side by cilia.

The eggs are oblique oval from a hundred measurements taken

from twenty five specimens, four from each, sixty per cent gave

25 u for length and 15.6 u for width. In ninety-fikve per cent

the maximum length was twebty-eight u and the remaining five per

cent twenty-seven u, the rest 23.4 u the minimum length. The

minimum width is 13 u and a width of 15.6 u. The width is more

constant than the length. Laurer's canal from the upper dorsal

side of the receptaculum seminie curves to the left side, takes

a posteriorly cou»ee dorsally over the left lobe of the ovary.,

0.18 mm below the receptaculum seminis. From there it passes

from a dorsal to a ventral course by that making a curve anterior

to the exoretory bladder. Then it makes a slight curve and opens

on the dorsal surface 0.054 mm from the excretory bladder. The

diameter of the opening is 0.024 mm. The longest diameter 0.04^

- 51 -

mm at the turning point from the ventral to dorsal surface. The

distance from the origin to the dorsal opening is about C.3 mm

but the actual length is much larger as it curves much in its

course. The walls of the canal is surrounded by circular muefcles

and cilia on the upper inner portion. At the opening to the out-

side there are glands opening into the canal. These appear to

be degenerate prostate glands. Tha* tends to show that Laurer's

canal is a vestigial vagina.

Seminal receptacle is pear or retort shape. The length

varies from 0.3 to 0.6 rum, the width from 0.18 to 0.26 mm, while

the thickness varies from 0.2 to 0.27 mm. The receptaculum seminis

lands from a median position toward the right side of the body in

few specimens on the left and in one median. The anterior end is

between the ovary lobes, the posterior part of it behind the

ovary. The seminal receptacle is somewhat more toward the dorsal

surface. The walls are non-nucleated . The sac-like cavity is

filled with spermatozoa. The anterior end of the receptaculum

seminin is tapering and aery narrow aboutO.018 mm. The Laurer's

canal makes a junction at the anterior end. From this junction

0.06 mm on the left side is the vitelline reservoir. The ovi-

duct opens just anterior to the Laurer's canal junction.

The uterus coils above anteriorly and to the right from the

ootype and curves dorsally from this point to the ootype. There

are no eggs in this portion which is however filled with sperma-

tozoa. The uterus above the ootype is 0.024 mm in diameter. The

uterine coils occupy nearly the middle 1/3 of the body. They are

filledwith yellowish eggs. The coils do not extend over the

intestinal crura. The number of ooils in different individuals

- 52 -

does not differ; the difference lies in the extent and the mag-

nitude of the coils. In the logg expanded individuals the coils,

are looser but in smaller and more contracted forms much more

compact. From the metraterm the uterus prolongs dorsally and

anteriorly to the left of the acetabulum at the lower margin of

which it curves ventrally. It extends 0.036 mm posteriorly and

coils dorsally and proceeds toward the right traneversly making

a dorso-ventral loop. From there it bends ventrally then dorsally

passing transversly between the two crura which I call the first

group. The margins of these loops are 0.12 mm from the left cms

and 0.06 mm from the right respectively. The second group extends

more toward the lateral crura, the left margin 0.18 mm, the right

0.036 mm. ^his group is formed by a transverse loop formed by the

coiling of the iterus toward the left ventrally and toward the

right dorsally. There are twenty- three suoh groups formed by

the coiling of the uterus do*so-ventrally and transversely.

Some of them are short and medium in size while others extend

from one crus to the other.

The shell glands are single isolated cells scattered in the

parenchyma anterior and anteeo-;ateral to the ovary extending up

to the uterus. They open into toke ootype. Their secretions are

known to be at present not shell material but yolk. Jhe glands

start nearly at the same plane as the ovary which is more ventral

The ecfccretory system - The excretory bladder occupies the

posterior fourth of the body on an average varying from 1/3.3

to 1/4.6 . The general shape is that of an S. The testes are

located in the concave spaces. The bladder ie well developed and

occupies almost the dorso-ventral space except at the excretory

- 53 -

pore region, and the testes region 2/3 to 4/5 of the body thickness

is occupied by the. bladder. This must vary as in preserved spec-

imens in most cases the exoretory bladder is greatly contracted.

Below the posterior testis the excretory bladder increases greatly

in diameter beooming a circular bulb about 0.38 mm in diameter.

There it narrows into a slit of 0.009 mm at the opening to the

outside. The pore is pesterior at the blunt tapering end of the

body, the length of narrow slit is 0.05 to 0.07 mm. The shape

of the excretory bladder in larval forms is straight but as the

growing proceeds adjacent otgans, the testes, expand and push

the bladder aside to make more room for themselves causing it tc

take an S shape, the present form as was suggested by Ward. The

forking of the anterior end in the genus Opisthorchis might be

due to the same cause , that is to the growth of the ovary post-

eriorly.

There are three kinds of excretory ducts, two main ducts,

one on each side of the body which carry waste material to the

excretory bladder. One seoondary duct unites with the main

duct at the level behind the acetabulum the other at the front

of the ovarial region. A branch comes frcim the posterior region

of the body. A third kind of branches unite to the main and the

secondary. These tubules terminate in flame dells. On account

of the poorly preserved specimens I have not been able to follow

every tubule. I have been able to see the flame cells in section.

They measure 1 to 1.1 micra. The ducts and tubules are symmetri-

cal on both sides of the body. The right main duct opens into

the excretory bladder anterior to the posterior testis and the

left 0.32 mm above the right branch. The main ducts have a very

-54-

wavy course between the intestinal crura and vitelline glands.

The alimentary system - The opening leading from the oral

sucker into the toharynx measures 0.04 to 0.5? mm in diameter.

The pharynx becomes somewhat larger in diameter toward the outer

margin of the sucker. The oral opening is terminal, continued

inside by the thinner cuticula. Behind the oral sucker there is

a little poekit 0.048 to 0.076 mm in diameter lined by a thin

lameiia. The pocket is somewhat funnel shaped. Behind the

pocket is more elliptical than the circular pharynx. The walls

of the pharyna are much thinner dorso-ventrally than laterally

so the pharynx has a verticle Bfcit appearance when focused

upon in the dor»o-ventral position. The thickness of the pharynx

is the same as the width. The inner lining of the pharynx consists

of thin cuticula 0.0013 to 0.0026 mm in thickness. The opening

of the oesophagus to the pharynx is much wider, about 0.047 mm

varying from 0.036 to 0.092 mm. The owsophagus runs 0.03 mm

ventrally to the middle line. It gradually increases in thickness

and width 0.057 mm. The anterior end ds circular gradually tak-

ing an elliptical form, the major axds being right to left. The

average width of the bifurcation is 0.19 mm. These figures

vary greatly according to the degree of contraction of the speci-

men at the time cf fixation. The owsophagus has an inner homogen-

eous layer varying in thickness from 0.006 to 0.012 mm. The layer

is surrounded on the outside by circular and longitudinal fibers.

There is no columnar epithelial cells at the bifurcation and the

oesophagus regions. The intestinal crura are lined with columnar

epithelial cells which are embedded in narrow elongatecells with

nuclei

.

- 55 -

The crura prolong posteriorly at first more median from

dorso-lateral surfaoe and gradually bend to the dorsal 0.06 to

0.08 mm. They curve in their course right to left more than

dorso-ventrally , but the later ones are greater in extent. It does

not overtop the uterine coils but takes a course between the

later and vi telling glands. It forms a trough in front of the

testes and a crest above. The same hapens ;at the reoeptauulum

seminis due to the gtowth of the these organs they push the caeca

to one side. At the posterior end the cmira bend slightly toward

the median line of the body which is occupied by the excretory

bladder. The crura in cross section are seen to be circular at It he

anterior region and somewhat irregular at the Altering and posterior

regions. This con be expected as the uterus fills with eggs and

consequently pushes the crura to one side, while the anterior

region will be left normal. The crura end blindly nearly to the

posterior end of the body. Generally both branches eiid at the

same level 0.21 to 0.4 mm from the posterior end of the body

0.25 mm being the average, sometimes the right or left extends

slightly further. The length of the crura if straight w6uld reach

to the posterior end. The crura are always full of blood which

shows that the parasite lives upon the blood of the host.

The cuticula - The body is covered by a non nucleated layer

the thickness of which varies in different regions of the body .

It being thicker at the oral end and around the ventral sucker .

The pharynx, the oral sucker, the ventral acetabulum, the genital

and the excretory pores are lined with a thin cuticular layer.

A portion of the cuticula stains more intensely than the other,

the deeper stained portion with Erlich's acid hematoxylin is the

- 56 -

outside layer and the lesser stained portion inside called by

some the basement membrane. There are four different kinds of

opinions as regards the origin of the cuticula. I would not at-

tempt to bring all their arguments but will only mention them.

The coverings of the invertebrates is in general a true

cuticii*. Some authors think Tremaftodes and Cestodes have a

cuticula seosreted by the hopodermis which means morphologically

that it is a true cuticula(Blochmann 1896). According to this

theory epithelial cells are sunken into the parenchyma and con-

nected to the cuticula by long projections.

A second si«w was that of Brandes (1892). He thinks the

outicula is secreted by the subcuticular single cells derived froim

the parenchyma and joined by ducts to the cuticula passing between

the superficial fibers. Tennent (1906) adopted Brandes view.

1 third theory is that epithelium (ectoderm) metamorphosed

into cuticula. Wagner (1855). Some of the authors supporting this

view are Monticelli, Goto, Nickereon and once by Braun. Sohwara

working with cercariae and Zeller with Polystomum notioed that

epithelial cells have their nuclei merely shrivelled.

Another theory originated by Leuckart (1886; 367) is that

after the disappearance of the epithelium the parenohyma portions

solidify in the peripheral regions by secretion of the cuticular

substance from the whole body of the parenchyma. Loose (1893,

1894) Braun (1893:818) Bratt (1898,1899) and Young (1908) confirm

this in there observations. Those who hold this view have var-

ious explanations to the possible function of the subcuticular

cells. Pratt suggested they secret some kind of protective sub-

stance against injurious products of the parasite. Looss and

- 57 -

Leuckart regard them aa connective tissue structures which give

rise to parenchyma cells. Loose compares it to the cambium of

plants. By my study of this form I am convinced that the cuti-

cula is formed as described by Young (1908:193) who worked with

Cyetocercus pdaiformis. His view is identical to that of Leucl-

art. Quoting him "The cuticula of Cyeticercue pisiformis is

developed from a ground of simple parenchyma fibrillae by a de-

position among them of a cement substance. There are bo spec-

ialized fibrillae or cellular processes concerned in its develop-

ment". I have seen those simple parenchyma fibrillae in the

cuticula in different parts of the same section. I have seen it

to be a continuation of the parenchyma fibrillae.

The parenchyma is formed of this strands of connective tissue

enclosing large, vacuolated spaces. Between the fibrous net work

there are small nuclei. All the body cavity is filled with these

parenchyma cells. The parenchyma cells near the circular muscles

are muoh smaller. It is the cortical layer of the eoto-parenchy-

ma of Brandes (1893)

In the parenchyma there are numerous giant cells few in the

pharynx and sucker muscles. These cells have many branches at

each side two main branches coming from the central portion

having the form of an epithelial cell nucleus. Each branch

divides into main branches and extends to the longitudinal mus-

cles, these cells are Nicoll*s myoblast, ganglion cells of Looss.

There are few of the same kind of cells as myoblast except the

various branches radiate from the common cinter and each radiat-

ing branch divides into fine fibers. They also extend to the

longitudinal muscles. These are called giant parenchyma cells.

In the form I have studied the subcuticular dells come

next to the longitudinal muscles in the parenchyma. They are

not arranged in a regular manner but are in a specific layer

which even in the totomounts the subcuticular cell layer is well

marked out. These cells have nuclei very much resembling the

parenchyma cell nuclei but it seems to me this cell has a gland-

ular nature rather than that of connective tissue structure gisiing

use to parenchyma cells. Pratt (1909 ) thinks these cells have

no ducts but are anastomosing branches which are interpreted as

ducts. In my form I have seen very many spindle shaped cells

their narrow ends toward the cuticula with a main branch which

extends to the cuticula. Whether these branches aee ducts or ncfct

I cannot definitely say, but I am inclined to think that they are,

as these branches are continuations from the cell which in some

cases clearly can be seen. But if these bells are glandular, what

is their function? Do they secret the cuticula as Blochman and

others think? Pratt's arguments against that view aae conclusive

enough. Cuticula is found in every parasitic form but some

ectoparasites have not the subcuticular glands. As there is no

definite experimental evidence to prove the nature of these cells

there is need of such work to prove their nature. They might

secret eome kind of enzyme to assist the cuticula in counteracting

the injurious substances existing in the host body.

Musculature - The muscles are fairly well developed.. Espec-

ially the muscles of the oral sucker and acetabulum. The pharynx

muscles are not as many as the sucker muscles. The suckers and

the pharynx have three kinds of muscles; radial, circular and

meridianal or longitudinal, the radial muscles are the strongest.

- 59 -

In the body there is another additional kind of muscle the

diagonal muscles passing from the right to the left or referse.

The muscles consist of bundles of very fine fibers in intimate

8Dttaot with each other. In a sagittal section, the circular mus-

oles are next to the basement membrane in a regular rectangular

block. In cross sections these are hemispherical. Next to the

circular muscles comes the longitudinal muscles. The diagonal

muscles can be seen in the parenchuma and are much finer . The

heaviest radial muscles pass dorso-ventrally . There can be seen

in cross sections muscles extending from the acetabulum to the

circular muscles called anchor muscles. The genital pore, the

excretory pore and the oesophagus are surrounded by ctrouiAr

and longitudinal muscles.

The nervous system - The nervous system in this form is

poorly developed. There is a central nervous system immediately

posterior to the pharynx. It is composed of two ganglia on each

side of the anterior end of the oesophagus united by a transverse

e

cpmmissure dorsally to the oesophagus. There are three branches

running anteriorly, the inner dorsal^ a median, and a ventral one

to the oral sucker, an exterior and a lateral. Three nerves

run backward from each ganglion, dorsal, ventral and lateral.

I have not been able to work out the complete nervous system

but it seems it does not differ from other species of this

genus

.

- 60 -

DISCUSSION.

Diagnosis as given by Leiper (1915:573-575)

Those worms resemble 0. felineus but differ in the following

points of specific importance. The skin is covered with minute

acicular spinelets. The ovary is multilobate. The branches of

the gut proceed almost to the extreme posterior end of the body.

The ovary and the testes fill the last fourth of the body between

the gut branches. The length of the oesophagus is from two to

three times that of the pharynx, varying with the degree of con-

traction of the specimens. The ventral sucker is about equal in

size to the oral, if anything occasioaally slightly larger.

Host - Indian civit cat, and man.

"In Dr. Kerr* e consignment of parasites obtained from prison-

ers in the Chiengmai jail was a tube containing three flukes re-

garding which the following notes were supplied" " Opisthorchis

felineus? ova found in about fifteen percent of the faeces examin-

ed, The worms were twice obtained post mortem, in one case a

single one from the intestine in another twelve worms from the

gall bladder and large bile ducts. The ova have a small projection

at the di3tal end, the cap. The worms also differ from the fig-

ures & havs ©f 0. felineus in having much larger part of the

intestine unbranched, they also seem smaller, I was present at

a post mortem by Dr. Mason where thousands of these worms were

removed from the liver."

From the above quotation and the previous diagnosis of the

form by Leiper who seems to have labeled the identif icatdonof the

form as 0. viverrini. My observations somewhat more in detail

confirms Leiper f s view. The form described in previous pages

- 61 -

corresponds to that of 0. viverrini. A comparative study of 0.

felineus and 0. viverrini will confirm this view.

0. felineus is muoh larger than 0.viverrini; the former

being from 8 to 13 mm the later 6mm. The form in question is

2.7 to 7*7 mm an average of 5.3 mm, corresponding to the later

one as the 2.7 forms were very much contracted very many of the

forms measured from 5 to 6 mm. The size of a form is a specific

character. The view that the size of a parasite has variations

with the size of the host is not accepted by prominent parasitolo-

gists. Looss, Braun and others. Loose (1907:42) afeainst the

view of Kat8urado stated that every species of parasite has a

size of its own. The occasional variation above normal is a

case just as is found in the higher animals.

The next most striking difference between 0. felineus and

0. vuverrini is in the size and shape of the ovary. In 0. fel-

ineus the ovary is small simple or slightly lobed in 0. viverrin i

it is larger and always lobed in the form under discussion, the

seminal receptaculum in 0. felineus is much larger compared to

tfte ovary than in 0. viverrini. The receptaculum seminis is

less prominent than ia 0. felineus and relatively smaller than

the ovary. Leiper in his paper reports the skin covered with

minute acicular spinelets; although I have examined numerous

specimens both toto mounts and sections, I have not been able

to see any spines. I have noticed in the outicula, or integu-

ment of some authors, deeply staining strands which might be

looked upon as small spinelets but more careful study of a

marginal sagittal section shows it to be of the same material as

the walls of the parenohyma that is of connective tissue.

-Ga-

in 0. viverrini Poiriar the suckers are small and equal0.28

rnra in the varl Chiengmai nearly same as the above 0.19 by 0.224

mm the oral and 0.226 by 0.23 mm the ventral on the other hand

O.felineus has 0.26 to 0.28 mm the oral sucker width and 0.228 mm

the length, the acetabulum 0.247 to 0.228 mm long and 0.197 to

0.228 mm wide. The pharynx in 0. felineus 0.204 to 0.169 tp

O.lBfl) mm to 0.170 mm in 0. viverrini 0.14 by 0.13 mm varying

from 0.13 to 0.18 mm in length and width 0.13 mm constant. An-

other marked difference between 0,felineus and the two kinds of

0. viverrini is the nelative length of the oesophagus. In 0.

felineus it is very short nearly ehe size of the pharynx but in

both varieties of 0. viverrini the oesophagus is about three times

as long as the pharynx.

The uterine coils in felineus are more compact and the

ooils do not correspond to that of 0. viverrini. But one of the

greatest evidences of these forms in wuestion not being an 0.

felineus is in the difference of the eggs of the two forms. The

two 0. viverrini are exactly alike, though my average estimate

for the 0. viverrini are exactly alike, though my average estimate

for the 0. viverrini var. Chiengmai is a little greater than

that given by Poirier. Poirier's form has a size 0.026 by

0.13 mm mine 0.026 by 0.0156 mm. Minimum size being 0.013 mm

In felineus the egg length varies 0.026 to 0.030 mm and the

width 0.011 to the 0.015 mm with an average of 0.030 by 0.011 mm

That is in 0. felineus the eggs are elongated and narrower, they

have smooth outer operculum with the shell, but in both 0.

viverrini the shell at the operculum takes a flask neck appear-

anoe instead of a smooth round ending as in 0. felineus.

- 63 -

In 0. felineus the exoretory system is Y-shaped in 0. viv-

errini Poirier not described but from the picture it is Host

likely similar to the new variety, which is sigmoid. The primary

ducts arising from different levels as represented in those figur-

ed .

Contrary to Looss's (1907:147) statement regarding the ex-

cretory bladder in the genus OPisthorchis being Jf-ahaped as one

of the differentiating characters of Clonorchis from Opisthorchis

the exoretory bladder in this form is like that of Clonorchis.

Having in mind the different preserving methods and change

in size of the forms accordingly in certain limits at certain

parts expeoially I conclude that the difference between 0. felin-

eus and this form is distinct enough not to diagnose it as a

new variety of 0. felineus but one of 0. viverrini Poirier.

Some of the characters described above are constant enough to be

regarded as specific peoularities . let as Braun, Luhe, Mtthling,

and Looss when studying the forms of the genus Opisthorchis have

spoken off the difficulty of distinguishing these species from

eaoh other, the same can be said for this new variety- In

anatomical detail, the topography of 0. felineus and this form

have very much in common. In fact these for^s have acicular spine-

lets as Leiper seems to have seen one cannot say it is not .

viverrini as Poirier in his description does not give any detail

in regard to thfct. Yet there stands a fact which might keep the

identity of these forms. Two forms might have identical features

but by different spedies on biological ground, that is in the

relation of the parasite to the hoet(Looss 1907:138). We know

that 0. felineus inhabits the cat, dog, glutton, fox and man

- 64 -

but 0. viverrini has not been reported before, aa being a normal

parasite of man. Poirier found the form from Felix viverinius

in France if the form found from Siam in identical to that found

by him toe might wxpect human ingection of the same kind in France

which ia not the case. The possible explanation in this case

would have bean simple if we could trace the history of the cat.

But as no one has reported O.viverrini after that we might assume

that it was an isolated case probably brought from other regions .

Supposing such an introduction from India suggests that

there must be cases in which dogs and cats in India are infected

which until now has not been reported as far as I am able to as-

certain. There is no doubt from the circumstances inwhich these

forms were secured that these are normal human parasites. But

the similarity of the forms with 0. viverrini Poirier and the

possible case of being a normal parasite in cats too gives some

certainty of its identity with 0. viverrini Poirier. Thru the

kindness of Dr. H. B. Ward I examined some of his collection of

0. felineus. One form from the liver of man collected by Dr.

Askanezy at Geneve, the other from St. Petersburg extracted by

Kowalewsky from the cat. The two forms have some miner variations

The form living innnan is muc& closer to the form in question

than that from the cat, the sides run parallel and greatest width

1.1 mm but the form from the cat was much broader 2.9 mm. The

oesophagus length 0.28 mm in the first ofte and <inly 0.228 mm in

the latter. From this comparison I can see the explanation of

the slight differsnces found between that of Poirier and mine.

From Poirier* s figure the form has not parallel sides but tapering

at the anterior end gradually widening till the posterior end

- 65 -

which narrows and ends as in the variety described. In the

figure given by Poirier the genitel pore is placed much farther

above the acetabulum than in forms I have examined. In his des-

cription Poirier states that the genital pore lies immediately

anterior to the acetabulum to I regard the representation in the

figure much farther than in fact it is. The posterior testis

is described and figured as four lobed, but as Poirier does not

mention how many forms he has examined I am inclined to think in

the normal case it has five. Rcom 50 individuals examined espec-

ially for this difference I found five with four lobes in many

cases one fcf the lobes poorly developed. The position of the

genital pore is constant and is specific but not the lobes of

the testis. In the typical fowns the anterior is four and the

posterior five lobes but this varies within certain limits. In

Leiper*s figure the lobes in the posterior testis are much

deeper than those I find in any onp of my specimens otherwise

the form is similar to mane except in none of the forms I have

studied do I find any spines as Leiper stated to have found in

his.

I add a few more records which might be of significance to

explaAA the possible occurance of 0. viverrini in man.

In 1907 Verdun and Bruyant recorded the occurance of 0.

felineus in a patient f*om Hanoft. The skin is stated to be

smooth and the ovary round or arcuate. Harwell sent from Formosa

a macerated specimen from the stool of a cat to Leiper which

had short spines on the cuticula. As the testes were digested

it could not be decided whether it is an Opisthorchis or Clonorchis

Looss (1907:152) reports a form sent to him by Chun which was

- 66 -

poorly preserved and could not be identified. But the form was

4.5 mm in length 0.9 mm in breadth. He calls attention to a

report from Ijima in 1886 which atates that the same kind of

forms naving short spine* in the skin were taken from the liver

of a cat. This form was regarded as a different form of Clonorchis

endemicus. Accordingly Looss said it might be another species.

Leiper thinks it might be 0. viverrini variety in question. One

thing definite is this that 0. viverrini Poirier is identical

with the form in question and anatomioally aas very close

resemblances to 0. felineus. From a biological standpoint there

is more definite proof this being a variety of 0. felineus which

is a normal parasite of man and also of the cat, that 0. viverrini

Poirier, which is reported definitely only from the cat. If it

can be proven that 0. viverrini var. Chiengmai in not parasitic

in the cat then it will become a new species instead of a

variety of 0. viverrini. But from the data at hand I can with

a certain degree of certainty say that tikis form is Opisthorchis

viverrini

.

But this only possible reasoning is less important if we

bear in mind that Feline Opisthorchis may occassionally occur in

man as stated by Looss (1907:153). Not only is this true in the

case fo Opisthorchis but in other forms as Paragonimus westermanai

and kellicotti anre found both in the dog and in man, in Japan.

Certain canine and feline parasites as Coccicium bigeminum,

Ascaris, Dipylidium are parasites in man also. Therefore I

diagnose this form as 0. viverrini Poirier var. Chiengmai, a

normal parasite fo the Indian civet cat not recorded heretofore

from man. In finishing I would express my thanks to Professor

H. B. Ward for hj^yaluable suggestions during this work.|

- 67 -

SUMMARY

1. The history of the genus Opisthorchis is given.

2. Speoifio diagnosis of the forms belonging to the genara

Opistorchis, Amphimerus and Paropisthorchis is given

with their generic diagnosis.

8 3. The relation of the speoies, the difficulty of their

identification is discussed briefly.

4. The description of a human parasite belonging to the

genus Opisthorchis is worked somewhat in detail and

diagnosed as Opisthorchis viverrini., var. Chiengraai.

- 68

LITERATURE

Aekanazy, Max

1900 Ueber Infektior. des Menechen mit Dietommm felineum

(Sibiricum ) in Ostpreussen und ihren zusammen-

hang mit Legerkrebs. Ctrbl Eakt. Paras, 1, 28:

491-5B2

.

1900a Distomum felineum in Ostpreussen beim Menschen.

Ctrbl. allg. Path. Path. Anat, 11:711-712.

1904 Die Aetiology tind pathologie der Katzenegelerkrank-

ung des Menechen. Beutech. Med. Wochenschr.

30:689-691

(0. felineus)

1906. Weitere Mitteilungen uber die quelle der Infektion

mit Distomum felinuem. Schrift. Phye. Gkon.

Ges. Kfinigsberg, 46:137-131.

(0. felineus)

Blanchard, P.

1895 An imaurn parasites. Bull Soo. Zool . France 20:217

(Opiethorchis

1895a Maladies paraeitamres , parasites animaux, para-

sites vigetaux a l'exclueion des Bacteries.

Traite de pathol . gen. de ch. Bouchard 8:649 -932

(Opiethorchis)

1895b Notices sur les parasites de l'homme. Ctrbl.

B&kt. u. Paras. 1, 17:2-13.

1900 Du role des ceux et des legumes dans l'etiologie

de 1 'helmSnthiase intestinale. Arch Paras.

- 69 -

3:485-491

.

(0. felineus)

Bossuat, Emile.

1902. Lea helminthes dans le foie. Arch, parasit.

5:161-206.

(0. felineua)

Braun, Max.

1893. liber die Distomum in der Leber der Hauakatzen.

Zool. Anz. 16:347-355.

(0. felineus)

1893. a Die Leberdistomen der Hauskatze (Felis catue

domesticua und verwandteArten) Ctrbl . Bakt. u.

Parae. 1 # 14:381-392, 422-428.

(0. felineus)

1894, Distomum Sibiricum. Bool. Anz. 17:128-129

1894a, Uber ein fur den Menschen neuee Diatomum aus der

Leber. Ctrbl. Bakt. u. Paras. 1, ;5: 602-606.

(0. felineus)

1900.. Ueber Campula oblonga Cobb. Ctrbl. Bak^. u. Parae.

1,28:8-9,249-252.

1901. Trematoden der Chelonier Mitt a, d. Zool. Mus.

Berl. , 2:34

.

(0. sinensis, 0. geminue)

1901a. Zur Revieion der Trematoden der Vogel. Ctrbl.

Bakt. Parae. 1, 29:560-568.

1902. Fascioliden der Vflfeel. Zool. Jahrb. Syst. 16:5-6

(0. inter ruptus, 0. lancea, Opisfck.)

1908. Die tierischen parasiten des Menschen. Ed. 4 :

186-190 (0. fel., O.nov., O.pseudo.)

- 70 -

Ciurea,

1914. Recherches Sur la Source de 1 1 infecticnd©

1 'homme et dee animauz par lee dietomee de la

famille dee Opiethorchiiaee . Bull. Beet. Sient.

Acad Roumaine Ann. 31:201-205

Cobbofcd, T. S.

1859. Synopeie of the Pietomidae. Journ. Lenn. Soc.

London, Zool . 5:1-56.

(O.nov.^ D. ccBjfcnctum)

1861. Further obeervation on entozoa with experimente

.

Trane. Linn. Soo. London. 23:349.

(O.nov.- D, conjuncture)

Cholodkoveky, N

,

1897. Dietomum felineuro in der Leber des Menechen.

Trav. Soc. Nat. St. Petereburg. 27:185-186.

(0. felineue)

1898. Ioonee helminthum hominie. 2 St. Petereburg.

(0. felineue)

1898a, Sur quelquee raree parasitee de lftHomme en Kuaado.

Arch. Paraeit. 1:354-355.

(0. felineue)

Dieaing, K. M.

1850. Syetema helminthum 1:334

<$0. lancea)

Engler, Kurt.

1904. Abnormer Darmverlauf bei 0. felineue. Zool. Anz.

28:186-188.

• 1

I

- 71 -

Galli-Valeria, Bruno.

1898. 0. pianae Nov. sp . eine veue Diatom idenart der

Wildente. Ctrbl . Bakt. 1;23:145-146.

1898a. Ueber Opisthorchis pianae fi.ep. Eine Erwiederung

an Herrn Prof. M. Kowalewski. Ctrbl. Eftkt. u.

Paras. 1, 24: 933.

1898 b. 0. pianae nov sp . eine neue Distomidenart der

Wildente. Ctrbl. Bakt. 1,23:145-146.

1901. La collection de parasites du laboratoire d'hygitoe

it de parasitologic de l'universite de Lausanne.

Bull. Soc Vandoise d. so. Nat., Lausanne 37:364.

(0. felineus)

Guerrini - Guido'.

1913. Ueber einen bis jetzt anbekannten fall parasitarer

infektion (0. felineus) in der Leber einee

Kaniachena) . Zeitschr. Infectionskrankl . paras.

Krankh. Hyg. Haustiere. 64: 262-270.

Gurlt, E. F.

1831. Lehrbuch der pathologiechen Anatomie der Haue-

Saugethiere 1:373-375.

(0. feleneus)

Hymann, George.

1905. Neue Distomen aus Cheloniern. Zool . Jahrb., Syet.

22: 98.

(Opisthorchis)

Hallack, Johanne.

1902. Zur Kenntntss der sexuellen Amphitypie bei Dicro-

coeliinen, Ctrbl. Bakt. 1,32:867-869.

(0. fel . , O.spec.)

- 72 -

Jacoby, Sever ia.

1900. Beitrflge zur Kenntnie einiger Distomen. Arch.

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1900. Sur 1 'opisthorchis tenuicollie (Rud. Mfthl ) =

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.

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.

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r

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.

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- 80 -

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203.

( 0. fel.)

ac acetabulum

am anchor muscle

cgl cephalic glands

cm Circular muscle

dan antero-dorsaii nerve

dm diagonal muscle

dvm dorsoventral muscle

eb excretory toladder

eph epithelial cell

as oesophagus

gc gamglion commissure

gp genital pore

it intestinal crura

lc. Laurerss canal

- 81 -

VII. EXPLANATION ©F PLATES

.

Abbreviations

os oral sucker

p parenchyma

pgl prostate gland

pd pedicle

ph pharynx

pn parenchyms nucleus

pph prepharynx

rs receptaculum seminie

rsu receptaculum seminie uter-inum

eg shell gland

sgl salivary gland

t. anterior testis

ta posterior testis

ut uterus

lco Laurer's canal dorsal opening

lm longitudinal muscle

mm meridonal muscle

my myoblast

od oviduct

ov ovary

va matraterm

vd vasdegerene

vdt vitelline duct

vr vitelline reservoir

vtd. vitelline transverse duct

vid. vitelline unpaired duct.

- 82 -

PLATE I • - Fig. 1 Opisthorchid sibiricum( = 0. felineus)

magnified after Winogradoff from Braun. Fig. 2 Egg of 0. fel-

ineus. Big. 3. 0. felineua (Riv.) after Braun magnified. Fig. 4

0. geminus x 10.5 Loose Plate 24 fig. 2 From Anas penelope

.

F4g. 5. Egg of 0. geminus x 631. Big. 6 0. simulans x81 after

Loose 1899. Fig.B Egg of 0. simulans x 631 Fig . 8 obsequens

x 24 after Nicoll. Fig. 9 0. tenuicollis xl5 tffter Braun from

Barker. (1911 pi 20 fig. 4.) Fig 10 Egg of 0. tenuicollis

Fig. 11. 0. longiseiinus xlO after Stiles and Hassall from Barker.

Fig 12, 0. viverrini magnified after Leiper. Fig 13. 01 viv

erini x 17 after goirier. Fig. 14. Egg of 0. viverrini

Fig. 15. viverrini central female organs.

Fig. 16. 0. simulans central female organs.

P1ATE II. - Fig 17 0pisthorohi8 geminus xll from Anas

boschas Looss 1899. Fig. 18 0. geminus egg. Fig 19. geminus

female central organs. Fig. 20 Amphimerus pseudofelineus x7fc

lard ( Reference handbook of medioal Science Fig, 4772) Fig.

21. A. pseudofelineus Central female organs. Fig. 22. A. pweude-

felineus egg. Fig. 23 A. fseudofelinews spines. Fig 24. A.

pseudofelineus genital region. Fig 25. A. oval is xl6.5 After

Barker . F±g. 26. A. ovalis spines. Big. 27. A. ovalis egg.

Fig. 38. A. ovalis genital region. Fig 29. A. lancea Diesing

xl7. ventral view after Weski . Fig. 30. A. speciosus xl5 After

Stiles and Hassall from Barker. Big. 31. A. speciosus egg.

Fig. 32". A. interrupt^ (from Aloedo bicolor ) x20 After Braun.

Big. 33. A, noverce agter Leuokart, from Braun. Fig. 84. A.

noverca egg. Fig. 35 Parepisthorehis caninus xl3 from Barker

- 83 -

Fig. 36. P. canimi8 spines.

PLATE III. Fig 37 - Opisthorohis viverrini, var. Chiengmai

x37 ventral view. Fig 38. 0. viverrini x30 dorsal view. Fig.

39. Female central organs xl35 dorsal view. Fig. 40 Female

Central organs x 135 ventral view. Fig 41. Genital region x 100

dorsal view. Fig 42. eggs of 01 viverrini x840. Fig. 43. An egg

of viverrini x 800. Fig 44. A was model x50 of the make

reproductive system. Fig. 45. From was model of genltia atrium,

represent idnthe opening of vas deferens and netraterm into that

common atrium. Fig. 46. A cross section x 340 representing tee

tall epithelial cells and surrounding narrow elongated nucleated

cells

•

PLATE IV. Fig. 47 A cross section of the acetabulum x340

the anchor, the radial meridional and circular muscles of the s

sucker represented. Fig. 48. A saggital section through the

acetabulum representing the vas defenens and ductus e jacuaatorus

and muscles of the acetabulum. Fig. A saggital section thru the

ovary and receptaculum seminis representing the dorsal open-

ing of Laurer's canal the ootype in which open single isolated

shell glands. Fig. 50. A cross seotion of acetabulum.

BLATE V. Fig. 51 A saggital section thru the oral sucker

and the pharyne. x 100. Fig. 52. A cross seotion thru the pharyns

xlOO. Fig 53. Cephalid fclands x 840. FUg. 54. A cross seotion

posterior to pharynx. Fig 55. Vitalline glands in the parenchyma

x 840. Figl 56. A saggital section x 340 showing muacxes.

Fig. 57. The central nervous system x 1G0. Fig 58. The excretory

system in part.

PLATE I

PLATE III

PLATE IV

1

PLATE V

I

»

I

1

j

3 0112 086765242